Cerebellum, motor and cognitive functions: What are the ... · 1 Cerebellum, motor and cognitive...
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Cerebellum, motor and cognitive functions:
What are the common grounds?
Eyal Cohen, PhD
(CogniFiber, CEO and Co-Founder)
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Cerebellum – The “Little Brain”
The Cerebellum takes ~10% of the Brain in Volume
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Small but Hefty…
Over 50% of the Brain’s Neurons are in the Cerebellum!!
The Convoluted Cerebellar Cortex consists of most Cerebellar Volume
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“Classical” symptoms of Cerebellar dysfunction
• Hypotonia = loss of muscle tone
• Tremor: limbs, jaw, neck, larynx
• Ataxia = loss of motor coordination:
1. Postural instability, “drunken sailor” gait , sway, wide standing base
2. Walking: uncertain, asymmetric, irregular
3. Failure in execution of planned movements i.e. intentional tremor,
dysmetria (lack of precision) and dysarthria (speech slurring)
4. Deficits in eye movement control
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Cerebellar is Central in Adaptation
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Position
Velocity
Biceps
Triceps
BicepsTriceps
Classical Role in Muscle Timing and Coordination
Cooling => Reducing Neuronal Firing
Villis and Hore 1977
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❖ Lower Intelligence (Verbal)
❖ Lower visuospatial abilities
❖ Memory problems (i.e. working, procedural) and Dementia
❖ Emotional control problems, impulsiveness, aggression
❖ Reduced ability of strategy formation
❖ Psychosis, Schizophrenia (co-morbid with reduced volume)
“Non-Classical” symptoms of
Cerebellar dysfunction
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Vermis
Hemispheres
FlocculusTonsil
Dorsal/Posterior ViewVentral/Anterior View
General Structure of the Cerebellar Cortex
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Vermis
Anterior Lobe
Posterior Lobe
Primary
Fissure
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Posterior/Dorsal view of the Cerebellum
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Anterior Lobe
Posterior Lobe
Cerebellar
Peduncles
FlocculusVermis
Tonsil
4th Ventricle
Nodulus
Nodules + Flocculi = Flocculu-Nodular Lobe10
Anterior/Ventral view of the Cerebellum
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Vestibulo-Cerebellum
Spino-Cerebellum Cerebro-Cerebellum
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Medio-lateral gross partition of the Cerebellum
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The 3D Folding of the Cerebellar Lobes
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Cerebellar Lobule
Cerebellar Folium
Cerebellar Folia
Lobes and Lobules of the Cerebellar Cortex
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The Cerebellum is not Only Cortex…
Deep Cerebellar
Nuclei (DCN)
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The Deep Cerebellar Nuclei (DCN)
are the Output Relays of the Cerebellum
FastigialInterposed
(Emboliform+
Globose)
Dentate
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16The Vestibular Nucleus in the Brainstem is
Functionally Homologous to cerebellar nuclei.
Functional mapping of the Cerebellar nuclei
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Inferior Olive (IO)
Pontine Nuclei (PN)
MF = Mossy Fibers
(~90% via the PN)
CF = Climbing Fibers
(All via the IO)
Two Major Input Pathways Serve the Cerebellum
Pontine NucleiInferior Olive
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The IO receive low level motor and sensory inputs
Visual Inputs:
SC = Superior Coliculus
NOT = Nucleus of Optic Tract
Vestibular Inputs:
VN = Vestibular Nucleus
Motor Command:
RN = Red Nucleus
Somatosensory & Proprioceptive:
DCN = Dorsal Column Nucleus
Trigeminal & Spinal Chord
Additionally: Auditory InputsSugihara & Shinoda JNS 2004
Inferior Olive (IO)
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Pontine Nuclei Relay Cerebro-Coritcal Information
(Brodal 1978)
Non-Cortical Inputs:
1. Mamilary Body
2. Amygdala
3. Midbrain Nuclei
4. Spinal Inputs
Forebrain Areas That project to the Pontine Nuclei
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Cerebellar Peduncles: Input / Output Highways
Medial: Pontine Inputs and Commissure
Medial Peduncle
Inferior Peduncle
Superior Peduncle
Superior: Thalamus/Midbrain Inputs/Outputs
Inferior: Spinal/Medullary Inputs/Outputs
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Cerebellar Major Output PathwaysMotor
CTXCTX = Cortex
PM = Premotor
PAR = Parietal
PF = Prefrontal
RN = Red Nucleus
VL = Ventrolateral
Thalamus
DCN = Deep Cerebellar
Nuclei
RF = Reticular Formation
PM
CTX
PF
CTX
PAR
CTX
VL
DCN
RN
RF
Other Outputs:
Inferior Olive
Hippocampus
Amygdala
Septum
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Mapping of Cerebellar Cortex – Classic View
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Cerebellar Cortex Mapping is Fragmented
Voogd and Glickstein 1998
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Cerebellar Cortex:
The Beauty of Network Architecture
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Purkinje Cells: The most Elaborate Neurons
of the CNS
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The 3 Layers of the Cerebellar Cortex
Molecular Layer
Purkinje cell Layer
Granule cell Layer
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27 Apps & Garwicz NatRevNeu 2005
Cerebellar Cortex Consists of 5 types of Neurons
Inhibitory cells:
• Purkinje
• Golgi
• Basket
• Stelate
Excitatory Cells:
• Granule cells
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28Modified from Apps
& Garwicz 2005
Deep Cerebellar
Nuclei (DCN)
Cerebellar
Cortex
Excitation
Inhibition
Climbing Fiber
~2000 Synapses
Parallel Fibers
~200,000 Synapses
PC: The Principal Cell of Cerebellar Cortex
Purkinje Cells are the only neurons
projecting from the Cerebellar Cortex!!
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The Spatial Organization of Cerebellar Circuitry
Notice the perpendicular
relationship between Mossy
Fiber and Climbing Fiber
enervations!!
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Parasaggital Microzones of Cerebellar Cortex
Some Molecular Markers (i.e. Zebrin II) divide Purkinje Cells Populations
into parasaggital stripes.
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Parallel fibers are not a delay line: local
inhibition prevents signal propagation
Yarom and Cohen AnnNYAc 2003
Stimulation of parallel fibersStimulation of granular layer
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32Modified from Apps
& Garwicz
NatRevNeu 2005
Closing the Loop: The Cerebellar Module
Deep Cerebellar
Nuclei (DCN)
Inferior Olive (IO)
Cerebellar
Cortex
Excitation
Inhibition
Feedback Inhibition
Feedback
Dysinhibition
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The simplified Olivocerebellar Module
Deep Cerebellar
Nuclei (DCN)
Inferior Olive
(IO)
Cerebellar Cortex
Cerebellar
Outputs
Thalamus,
Red Nucleus,
Brainstem
Mossy Fibers
Sensorimotor
Context (forebrain)
Sensory Feedback
+ Efferent Copy of
Motor Command
Parallel Fibers
Climbing FibersPontine Nuclei
(PN)Purkinje Cell
Granular Layer
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Cerebellar Physiology
Double Recording of Purkinje Cell in Slice
Hausser M.
Infra Red Fluorescent Dye
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Purkinje Cells exhibit Two distinct Spike Types
Mossy Fiber Stimulation Inferior Olive Stimulation
Eccles, 1966
Simple Spike Complex Spike
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Cerebellar Learning Theories
Eccles, 1967
1971
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Simple Spikes are Modulated by Inputs
Roitman 2005
Simple spikes of 2 Purkinje cells in awake monkey during hand movements
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Olivary Spikes are fired in Unexpected Events
Horn, Pong & Gibson 2002
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Distinct Firing Regimes of Purkinje Cells
Complex Spikes : 1-3Hz (Phasic 10-15Hz)
Simple Spikes: 0 - ~80Hz
(Mostly 0Hz or 4-8Hz or 20-80Hz, with possible phasic >100Hz)
100ms
10mV
Complex Spikes & Simple Spikes (Up-State)
Complex Spikes Only (Down-State)
Cohen and Lamp (unpublished)
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Complex Spikes as Purkinje Cell Switches
Loewenstein et al. NatNS 2005
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Synchronous Population Coding by Purkinje Cells
Welsh 1995
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Saccade attributes are encoded by
a population of ~100 Purkinje cells
Tier-Barash 2000
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Vestibulo - Ocular Adaptation (VOR)
Ito 1984, Schonville et al. 2010
VG = Vestibular Ganglion
VN =Vestibular Nucleus
PA = Pontine Area
AOS = Accessory Optic
System
OM = Oculomotor
NeuronsRetinal Slip
Climbing
Fiber
ActivationMotor
Correction
Synaptic
Changes
Reduced
Retinal Slip
Visual Error
Complex
Spike
Synaptic
Changes
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Plasticity in the Cerebellar Cortex
Coesmans et al. Neuron 2004
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Eyelid Reflex Conditioning
Bracha and Bloedel 2009
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Eyelid Reflex Conditioning
Medina et al. Nature 2002
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Eyelid Reflex Conditioning
Rasmussen et al 2008
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IO is crucial for Learning @350ms but not 700ms
Welsh 2002
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The Learning Transfer Hypothesis
DCN
Cerebellar
CTX
Fastest Learning~
Sec-Min
VL = Ventro-Lateral Nucleus
DCN = Deep Cerebellar Nuclei
CTX = Cortex
CF = Climbing Fibers
Inferior Olive
Motor
CTX
VL
After 6 Hrs
After 12 Hrs
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❖ All physiological results so far:
• in animals
• related to motor function
• or coordination
• or timing
❖ Can timing be also related to higher cognitive functions?
❖ Is there evidence from human real-time physiology?
❖ Are there specific areas in the cerebellum which relate to cognitive
functions?
What about the “Non-Classical”
symptoms?
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Specific cerebellar regions with cognitive
dysfunction following focal damage
Cognitive effects of focal damage were found when:
1. The damage involved the vermis
2. The damage was in an areas which get blood supply from the posterior inferior
cerebellar artery
3. Children with Vermal damage show autistic-like features:
➢ Irritability
➢ impulsivity
➢ Disinhibition
➢ Emotional lability
4. Complex verbal dysfunction associated with right cerebellar damage
5. Dysprosodia (pronunciation fault): associated with left cerebellar damage
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Somatosensory Processing
FMRI Mapping pf Cerebellar Involvement in
Various Processes
Stoodley and Schmahmann 2009
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FMRI Mapping pf Cerebellar Involvement in
Various Processes
Motor Processing
Stoodley and Schmahmann 2009
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Language Processing
FMRI Mapping pf Cerebellar Involvement in
Various Processes
Stoodley and Schmahmann 2009
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Working Memory
FMRI Mapping pf Cerebellar Involvement in
Various Processes
Stoodley and Schmahmann 2009
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Spatial Processing
FMRI Mapping pf Cerebellar Involvement in
Various Processes
Stoodley and Schmahmann 2009
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Executive Processing
FMRI Mapping pf Cerebellar Involvement in
Various Processes
Stoodley and Schmahmann 2009
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Emotional Processing
FMRI Mapping pf Cerebellar Involvement in
Various Processes
Stoodley and Schmahmann 2009
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Cerebellar Involvement in Autism
Control Subject Autistic Subject
Levitt J. 1999
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Cerebellar Involvement in Dyslexia
Control Subject Dyslexic Subject
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Autism candidate genes affects the Cerebellum
Control CASP2 -/-Autistic Control
Lose of Purkinje Cells in Autistic Developmental Problems in KO mice
Saskia 2004 Sadakata 2007
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Olivocerebellar Information flow : Mossy Fibers Inputs
Cerebellar
CTX
Motor
CTX
Sensory &
Vestibular
Information
Pontine Nuclei
Motor Command
To Spinal MNs
VL = Ventro-Lateral Nucleus
DCN = Deep Cerebellar Nuclei
RN = Red Nucleus
(+ mid brain nuclei)
CTX = Cortex
MF = Mossy Fibers
CF = Climbing Fibers
PM
CTX
PF
CTX
PAR
CTX
VL
MF
RF
Afferent Copy
Of Motor Command
RN
DCN
Inferior Olive
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Cerebellar
CTX
Motor
CTX
Sensory &
Vestibular
Information
Pontine Nuclei
Motor Command
To Spinal MNs
VL = Ventro-Lateral Nucleus
DCN = Deep Cerebellar Nuclei
RN = Red Nucleus
(+ mid brain nuclei)
CTX = Cortex
MF = Mossy Fibers
CF = Climbing Fibers
PM
CTX
PF
CTX
PAR
CTX
VL
RF
Afferent Copy
Of Motor Command
RN
DCN
Inferior Olive
Olivocerebellar Information flow : Climbing Fibers Inputs
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Hypothesized information flow during Cerebellar function
Cerebellar
CTX
Motor
CTXVL = Ventro-Lateral Nucleus
DCN = Deep Cerebellar Nuclei
RN = Red Nucleus
(+ mid brain nuclei)
CTX = Cortex
MF = Mossy Fibers
CF = Climbing Fibers
Motor
Learning
PM
CTX
PF
CTX
PAR
CTX
VL
Motor Real-Time
Correction
RN
DCN
Inferior Olive
RF
Pontine Nuclei
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The General Role of the Cerebellum
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The Inferior Olive as the Great Comparator
Inferior Olive
Sensory
feedback
Motor Intention
Feedback Inhibition from
Deep Cerebellar Nuclei
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IO Neurons are Coupled by Gap Junction
Placantonakis et al. PNAS 2004
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The Inferior Olive as the Great Comparator
Inferior Olive
Sensory
feedback
Motor Intention
Feedback Inhibition From the DCN
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IO Neurons are Natural Oscillators
Llinas and Yarom JPhysiol 1986
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IO neurons are not Harmonious Oscillators in-vivo
Chorev, Yarom and Lampl JNS 2007
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There are Various Oscillation Patterns
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Oscillation as a Blocking Device for expected Inputs
Expected Inputs
How will The IO Differentiate Early vs. Late Inputs?
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73 Medina et al. Nature 2002
Conditioning Memory Retention
depends on Nucleo-Olivary Inhibition
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ISI vs. Oscillation phase “encoding”
Mathy, et al. 2009
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CF activation Pattern sets the direction of plasticity
Mathy, et al. 2009