Motor systems: Reflexes, pyramidal and extrapyramidal

system, cerebellum
Contents:

1. Muscle, muscle spindle, motor unit and

reflexes
2. Voluntary movement: Organization
and major pathways
3. Extrapyramidal system of movement
- Basal ganglia
- Cerebellum

Literature:
Dudel et al., Neurowissenschaft (Springer)
Reichert, Neurobiologie (Thieme)
Kandel et al., Principles of Neural Science (McGraw Hill)
Kahle, Taschenatlas der Anatomie, Band 3: Nervensystem und
Sinnesorgane (Thieme)
Greenstein and Greenstein, Color Atlas of Neuroscience (Thieme)
 Motor systems

Motor
Integration
Output
(Interneurons)
(muscles)
 Muscle, muscle spindle, motor unit and reflexes
Organization of muscles
Skeletal muscle
→ Muscle fiber (fused cells, surrounded
by a plasma-membrane (sarcolemma))
→ myofibril (surrounded by
sarcoplasmic reticulum)
→ sarcomere
(functional motor unit)

Contains thick and thin

filaments (myosin,
F-actin)

Z-disk

Sliding filament
hypothesis
(A.F. Huxley et al., 1950s)
1.5 – 3.5 µm
 Muscle, muscle spindle, motor unit and reflexes
Sensing muscle tension
Muscle spindles: encapsulated sensory receptors
located within muscles
signal changes in the length of the muscle (stretch sensor)

Anatomy: encapsulated muscle fibers located parallel to the muscle fibers

large-diameter sensory endings in the middle noncontractile portion
and small sensory endings at contractile portions (blue),
small-diameter motor endings at the polar portions (gamma motor
neurons; red)

(Kahle, Taschenatlas der Anatomie Bd. III)

 Muscle, muscle spindle, motor unit and reflexes
Basic motor unit: a-motoneuron in the
ventral horn + innervated muscle fibers
Most of efferent tracts (orange) do not directly end at
a-motoneurons but at interneurons (black) →
complex integration
Peripheral tracts (blue) may directly end at a-
motoneurons

(Kahle, Taschenatlas der Anatomie Bd. III)

 Muscle, muscle spindle, motor unit and reflexes
Basic motor unit: a-motoneuron in the ventral horn + innervated muscle
fibers

Strength Speed
Recruitment of motoneurons Tetanic contraction (physiologic tetanus):
Sustained muscle contraction evoked
when the motor nerve that innervates a
skeletal muscle emits action potentials at a
very high rate (interval < 75 ms)

(Campbell, Biologie)
 Muscle, muscle spindle, motor unit and reflexes

Coordinated work of muscles

Muscles pull but cannot push → hinge requires at least two antagonist muscles
 Muscle, muscle spindle, motor unit and reflexes
Reflex pathways
What are reflexes?
Involuntary coordinated patterns of muscle contraction and relaxation
elicited by peripheral stimuli
traditionally seen as automatic, stereotyped movement
modern view: can be modified via supraspinal signals

Test: complete transection of the spinal cord from the brain

Receptors in muscles: stretch reflexes

Cutaneous receptors: withdrawal reflexes
 Muscle, muscle spindle, motor unit and reflexes

Stretch reflex

Receptors in muscles (spindle)

Most studied and most simple

reflex: Contraction of a
muscle when the
muscle is lengthened

Experiment by Sherrington
(beginning of 20th century):
reflex was abolished by cutting
either dorsal or ventral root

Involves monosynaptic pathway

Heteronymous innervation

Reciprocal innervation
(Kandel, Principles of Neural Science)
 Muscle, muscle spindle, motor unit and reflexes

Flexion-withdrawal reflex

Cutaneous receptors

Polysynaptic pathway

Reciprocal innervation

Crossed-extension reflex
 Muscle, muscle spindle, motor unit and reflexes

Modification of reflexes

Supraspinal contacts to alpha or

gamma motoneurons

„Spinal shock“
Voluntary movement: Organization and major pathways
Control of subcortical motoric centers by the cortex

Sulcus centralis

4
6
Voluntary movement: Organization and major pathways

Control of subcortical motoric centers by the cortex

primary motor cortex (area 4) →

lowest intensity of stimulation
elicits movement
6 4
premotor cortex (area 6)

Motor homunculus
 Voluntary movement: Organization and major pathways
Major pathway of voluntary motor information: Corticospinal tract
(“Pyramidenbahn”)

Massive bundles of fibers (approx. 1 million of

axons)
Originates from primary motor cortex, premotor
cortex and somatosensory cortex

Somatotopic organization
Main tract crosses at the
medulla (pyramidal
decussation) and
descends in the lateral
column (Tractus
corticospinalis lateralis)
Most end at interneurons
between dorsal and
ventral horn
Pyramidal decussation
Voluntary movement: Organization and major pathways

Majority (70-90%) of the axonal fibres cross at the

pyramidal decussation
Remaining uncrossed fibers descend in the ventral
column (Tractus corticospinalis anterior) and cross
at the position of their endings

Tractus corticospinalis
lateralis

Tractus corticospinalis
Control more
anterior
distal
limb muscles
Control of posture by integrating →important for
visual, vestibular and goal-directed
somatosensory information movement
 Voluntary movement: Organization and major pathways

Corticospinal tract usually ends at Zona intermedia between ventral and

dorsal horn at interneurons

Minority directly contact motoneurons at the ventral horn (in most cases
motor neurons that innervate flexors of the distal limb) → direct control
by the corticospinal tract
 Extrapyramidal system of movement

Phylogenetically old extrapyramidal

motor system
multisynaptic chain of neurons
In the narrower sense: group of nuclei
with a high iron content (basal
ganglia)
In a broader sense: together with the
cerebellum as an integration center

Function: regulation of involuntary

movements for keeping posture and
trained movements
Acts as a „servo-mechanism“ for
voluntary movement and provide feed-
back loops
→ required for „smooth“ movement
 Extrapyramidal system of movement

Basal ganglia

Striatum (putamen and caudatum)

Pallidum
Nucleus subthalamicus
Nucleus ruber
Substantia nigra

Belong to different parts of the

brain:
telencephalon (Striatum),
diencephalon (Pallidum, N.
subthalamicus)
brain stem (N. ruber, S. nigra)
 Extrapyramidal system of movement

Basal ganglia
Highly connected with each other and the cortex and cerebellum

Pallidum Striatum (caudatum)

Striatum (putamen)

Nucleus
subthalamicus

Nucleus ruber Substantia nigra

Striatum: integration unit of the extrapyramidal motor system

Characterized by the presence of striasomes (patches) which stain for neuropeptides
and opioid receptors and a matrix which stains for acetylcholinesterase
 Extrapyramidal system of movement
Connections of the basal ganglia (schematic representation)
Nigrostriatal pathway

Reciprocal coupling between - Striatum and S. Nigra

- Pallidum and N. subthalamicus
 Extrapyramidal system of movement
Extrapyramidal syndromes
Lesions in the basal ganglia lead to characteristic disturbances of movement:

e.g., Lesion of the N. subthalamicus:

Hemiballism – involuntary (often violent) movement of the limbs

e.g., Degeneration of the Striatum:

Chorea Huntington – involuntary movement, cognitive impairment (dementia)
genetic disease

Parkinson‘s disease
Most frequent disease of the motoric system
(incidence: 1-5% of 70-90 year old
people)
paucity of spontaneous movement
(„Maskengesicht“), increased muscle
tone (rigidity), characteristic tremor at rest
(„Schüttellähmung“)
 Extrapyramidal system of movement
Cerebellum: Integration unit of the extrapyrimadal system

Latin: „little brain“, constitutes 10% of the brain volume but 50% of its neurons
Neurons are arranged in highly regular manner as repeating units → basic
circuit modules
Major output: premotor and motor cortex, basal ganglia of the brain stem
Many parallel convolutions called folia („leaves“)
Connected to the brain stem via pedunculi cerebellaris („Kleinhirnstiele“)

Functional structure:
 Extrapyramidal system of movement
Cells of the cerebellar cortex
Neurons are highly ordered and organized in repeat units → „modules“

Only five types of neurons:

- Stellate neurons (“Sternzellen”)
- Basket neurons (“Korbzellen”) Inhibitory
- Purkinje neurons neurons
- Golgi neurons
- Granule cells (“Körnerzellen”) Excitatory
neurons

(Fischer et al. (1998) Neuron 20: 847-854)

 Extrapyramidal system of movement

Organization of the cerebellar cortex

Organized in
3 layers:

Molecular layer: cell bodies of stellate and basket cells, axons of granule cells
(oriented as parallel fibers along the “folia”), dendrites of Purkinje cells
(oriented perpendicular to the parallel fibers)
Purkinje cell layer: single layer of Purkinje cell bodies, axons project to white
matter
Granule layer: many granule cells and some Golgi neurons
 Extrapyramidal system of movement
Input connections of the cerebellar cortex
Two main types of inputs:
- input from mossy fibers - input from climbing fibers
 Extrapyramidal system of movement
Connections within the cerebellar cortex
1. Axons of granule cells travel to
molecular layer and excite as
“parallel fibers” many Purkinje
neurons in the same transverse
plane

2. Basket cells and stellate cells

make inhibitory contacts to
Purkinje neurons, thus
producing an inhibitory side-
loop
 Extrapyramidal system of movement

Output connections of the cerebellar cortex

Axons of the Purkinje neurons project
into the white matter (deep nuclei of the
cerebellum) and provide the (entirely
inhibitory) output of the cerebellar
cortex (mediated by GABA)
 Extrapyramidal system of movement

Principles of function of the cerebellar cortex I

Incoming signals from mossy fibers
(from spinal cord and brain stem)
synapse at dendrites of granule and
golgi cells

Parallel fibers of granule cells excite

only one row of Purkinje cells

Golgi cells

Golgi cells are much larger than granule cells and

have dendrites filling a large volume in all directions →
are excited in a larger volume and locally inhibit (via
short axons) neighboring Purkinje cells

→ Enhancement of contrast
 Extrapyramidal system of movement
Principles of function of the cerebellar cortex II
Inhibitory interneurons

Neurons of the subcortical nuclei are

excited by axon collaterals of
climbing and mossy fibers but
inhibited by Purkinje neurons.

Thus, the output to the descending

motor systems is modulated by the
cortical side loop.

Neurons of
subcortical nuclei
Climbing fiber
Mossy fiber
 Extrapyramidal system of movement
Principles of function of the cerebellar cortex II

Neurons of the subcortical nuclei are

excited by axon collaterals of
climbing and mossy fibers but
inhibited by Purkinje neurons.

Thus, the output to the descending

motor systems is modulated by the
cortical side loop.
 Extrapyramidal system of movement
Principles of function of the cerebellar cortex III

Neighboring purkinje cells

receive signal of the
same granule cell via the
parallel fiber with
increasing delay
(speed of conductance of a
parallel fiber: 0.2 m/s;
corresponds to about 0.1
ms for every neighboring
Purkinje cell)

→ Temporal correlation of signals can be determined and movement

(activation of muscle fibers) can be segmented (there are often difficulties
with cerebellar lesions)
 Extrapyramidal system of movement
The cerebellum in motor learning
e.g. vestibulo-ocular reflex: ensures that the eyes can fix a target when turning the
head.
When wearing prismatic glasses, the reflex turns after a learning phase.
Destruction of the vestibulocerebellum prevents this adaptation
 Extrapyramidal system of movement
The cerebellum in motor learning
 Extrapyramidal system of movement
The cerebellum in motor learning
Simple model system: Rabbit eyelid conditioning

Conditioned Mossy fiber

tone
Stimulus (CS)
Cerebellar
cortex
Unconditioned Climbing fiber
puff
Stimulus (US)

Learning depends on the interstimulus interval (ISI):

Deep nuclei of the

cerebellum
(Interpositus nuclei)

(from: Ohyama et al. (2003) What

the cerebellum computes. Trends
Neurosci. 26: 222-227)
 Extrapyramidal system of movement
The cerebellum in motor learning
Simple model system: Rabbit eyelid conditioning
 Extrapyramidal system of movement
The cerebellum in motor learning
Simple model system: Rabbit eyelid conditioning

Cerebellar cortex
Mossy
Climbing
fiber
fiber
+ +
Puff (US) - (CS) tone

+ +

Deep nuclei of the

cerebellum
(Interpositus nuclei)
 Extrapyramidal system of movement
The cerebellum in motor learning
What is the mechanism of learning during eyelid conditioning?
Hypothesis:
- Changes of synaptic structure in the interpositus nuclei
- Changes would need to be relative long lasting to support long-term
encoding of the CS/US association
→ Formation of new synapses may be involved
In fact: Increase in the number
of synapses per neuron
 Extrapyramidal system of movement
The cerebellum in motor learning
Which type of synapse is
involved?

Excitatory synapse Inhibitory synapse

(from: Kleim et al.

(2002) Synapse
formation is
associated with
memory storage
in the cerebellum.
J. Neurosci. 99:
13228-13231)
 Extrapyramidal system of movement
The cerebellum in motor learning

Parallels between cerebellum-

and amygdala-dependant
conditioning
Javier F. Medina, J. Christopher
Repa, Michael D. Mauk &
Joseph E. LeDoux. Nature
Reviews Neuroscience 3, 122-
131(2002)