Advanced Neuroanatomy:

Basal Ganglia and Cerebellum

Lecture 5

NEURONproject
Dr. Mojgan HODAIE
Cerebellum
Cerebellum

• Latin for “little brain”, it is only

10% of the volume of the brain
but contains over 50% of total
neurons
• Located underneath the
cerebral cortex separated by the
cerebellar tentorium
• Divided along the midline by the
vermis
• Highly convoluted with folds
called folia
 Cerebellum
• The cerebellar cortex is divided into three lobes:
1. Anterior Lobe
2. Posterior Lobe
3. Flocculonodular Lobe
• Functionally divided into three sections:
1. Spinocerebellum
2. Cerebrocerebellum/Pontocerebellum
3. Vestibulocerebellum/Archicerebellum
• The Anterior and Posterior Lobes are separated by the
primary fissure
• The Posterior and Flocculonodular Lobes are separated by
the posterolateral fissure (also called posterior fissure)
 Cerebellum
• Important component of the central motor control system
• Integrating center for afferent sensory and other inputs,
receives information from:
– Ascending spinal tracts
– Cerebral Cortex
– Eye
– Vestibular Apparatus
• Provides major input to the brainstem and cortical pathways for
control of the limb, trunk and eye movement
– Regulates movement indirectly through adjustment of the output of the
major descending motor systems of the brain
• Influences speech and balance as well
 Spinocerebellum

Cerebrocerebellum/
Pontocerebellum

Vestibulocerebellum
 Spinocerebellum
• Receives somatic sensory input from the spinal
cord
• Also receives information from the trigeminal
nerve and other sensory cranial nerves
• Important role in regulating posture and
movements of the trunk and limbs
 Spinocerebellum
• Important control of the musculature of the body
• The Dorsal Spinocerebellar Tract originates from the
Clarke’s nucleus to provide sensory information
regarding the leg and lower trunk
• The Cuneocerebellar tracts originates from the
Accessory Cuneate nucleus to provide sensory
information regarding the arm and upper trunk
• Afferent signals from these tracts connect to the Red
nucleus (Magnocellular division), Ventrolateral nucleus
(Thalamus) and to motor areas in the Frontal lobe
– These later give rise to the lateral descending pathways for
motor control
 Cerebrocerebellum/Pontocerebellum

• Indirectly receives input from the cerebral

cortex
– Major input from the contralateral cerebral cortex
(motor, sensory and association areas)
• Important function in the planning of
movements and non-motor functions
 Cerebrocerebellum/Pontocerebellum
• The Cerebrocerebellum’s Purkinje neurons
project to the Dendate nucleus
– These neurons project to two major motor control
centres:
• Ventrolateral nucleus (Thalamus) to the Primary motor
cortex and premotor areas
• Red nucleus (Parvocellular division) to the ipsilateral
inferior olivary nucleus
• Has connections to areas that mediate higher
brain functions as well
 Vestibulocerebellum
• Directly receives input from the vestibular labyrinth and
vestibular nuclei
• Assists in maintaining balance as well as regulating head and
eye movements, important for controlling gaze
• The Vestibulocerebellum projects back to vestibular nuclei
(lateral, medial, inferior and superior)
– Coordinates neck muscles with eye control through the medial
vestibulospinal tract
– Maintain balance through the lateral vestibulospinal tract
– Regulate eye movement through the medial longitudinal fasciculus
 Functional Anatomy
• Cerebellum resides in the posterior cranial fossa
• Contains three pairs of deep nuclei:
– Fastigial
– Interposed (includes globose and emboliform nuclei)
– Dendate
• Important connections called peduncles
– Bundles of fibers that connect the cerebellum with the
brain stem
 Cerebellar Peduncles
• There are a total of six peduncles
• Appear in pairs:
1. Superior Cerebellar Peduncle (Brachium
Conjunctivum)
2. Middle Cerebellar Peduncle (Branchium Pontis)
3. Inferior Cerebellar Peduncle (Restiform Body)
Cerebellar Peduncles
 Superior Cerebellar Peduncle
• Conveys mainly efferents, is the major output pathway of the
cerebellum
– There are a few afferent fibers originating from the ventral
spinocerebellar tract that enter through this peduncle to the
anterior cerebellar lobe
• Dendate nucleus is origin of most efferent fibers, project to:
– Red nucleus
– Ventral lateral and ventral anterior nuclei of Thalamus
– Medulla
– Other midbrain structures
• Two major pathways involved in motor planning pass through
this peduncle
 Middle Cerebellar Peduncle
• Made of only afferent fibers from the pontine nuclei as part of
the corticopontocerebellar tract, includes many fibers from
sensory and motor areas which makes this peduncle the
largest
• Fibers are arranged into three fasciculi:
1. Superior
• Most superficial, mainly distributed to lobules on the inferior surface of
the cerebellar hemisphere
2. Inferior
• Underneath the superior fasciculus, distributed to the folia near the
vermis
3. Deep
• Covers the inferior peduncle, distributes to the upper anterior cerebellar
folia
 Inferior Cerebellar Peduncle
• Includes important fibers such as the dorsal spinocerebellar
tract and axons from the inferior olivary nucleus
• Mostly input and output fibers necessary for integrating
proprioceptive sensory input with motor vestibular function
• Posterior/dorsal spinocerebellar tract carrying proprioceptive
information passes here to synapse within the
spinocerebellum
– Vestibular information is provided to the vestibulocerebellum
• Cuneocerebellar tract passes through as well
• Also carries information from Purkinje cells out to the
vestibular nuclei of the brainstem
 Summary of Peduncles
Afferent Source Information Type Peduncles Involved

Ipsilateral vestibular Head position and Inferior cerebellar peduncle

apparatus acceleration

Ipsilateral proprioceptors Body position Superior and inferior

cerebellar peduncles

Ipsilateral superior Visual input Superior cerebellar

colliculus peduncle

Ipsilateral inferior colliculus Auditory input Superior cerebellar

peduncle

Contralateral cerebral Information about Middle cerebellar peduncle

hemisphere voluntary movement that is
being planned or ongoing
 Functional Anatomy
• Two major sets of input:
– Climbing fibers (originating from the Inferior Olivary Nucleus)
– Mossy fibers (originating from various brain stem and spinal cord
nuclei)
• Climbing fibers synapse on Purkinje neurons in the cortex
which send signals to the deep nuclei of the cerebellar cortex
• Mossy fibers interact with both excitatory and inhibitory
interneurons
– Excitatory ones synapse on Purkinje neurons leading to output
signals to the cerebellar cortex
– Inhibitory neurons have variable effect on Purkinje neurons
 Climbing Fibers
• Origin is the inferior olivary nucleus
• Conveys somatosensory, visual or cerebral cortical
information
• Closely wrapped around cell bodies and dendrites of the
Purkinje neurons making several connections
• Generates depolarization of Purkinje neurons through
prolonged voltage-gated calcium conductance which
causes a complex spike
– Complex spikes are a large initial amplitude spike followed by
a high number of smaller amplitude action potentials
 Mossy Fibers
• Originate from nuclei of spinal cord and brain
• Carry information from periphery and from cerebral
cortex
• Terminate as excitatory synapses on dendrites of
granule cells
• The long axons of granule cells (parallel fibers) are
able to excite the Purkinje neurons close to the
molecular layer
– Parallel fibers cause a single action potential or simple
spike
 Cortical Layers
• 3 layers comprise the cerebellar cortex,
from outermost to innermost they are:
1. Molecular layer
2. Purkinje layer
3. Granular layer
• Five types of neuronal cells exist within
the cerebellar cortical layers:
1. Inhibitory stellate
2. Basket
3. Purkinje
4. Golgi neurons
5. Excitatory granule cells
 Cortical Layers
• Molecular layer:
– Contains inhibitory interneurons: stellate and basket cells
– Lie amongst the excitatory axons of granule cells and dendrites of inhibitory
Purkinje cells that arise from other layers
• Purkinje cell layer:
– Single layer of Purkinje cell bodies
– Dendritic extensions of the Purkinje cells extend upward into the molecular layer
– Axons project to deep cerebellar and vestibular nuclei, providing the inhibitory
output of the cerebellar cortex
– Purkinje neuron activity is inhibited by stellate, basket and Golgi interneurons
• Granular layer
– Contains mainly granule cells and few Golgi interneurons
– Site of mossy fiber termination, they contact granule cells and Golgi neurons in
synaptic complexes called cerebellar glomeruli
 Motor System
(Cerebellum)
• The cerebellum doesn’t directly generate
movement but its crucial in coordinating
movements
• It appears to tonically excite motor centers
including brainstem and thalamus
• Fastigial nucleus: regulates posture and locomotion
• Interposed nuclei: regulates reach and grasp
• Dentate nuclei: regulates fine skills (such as writing)
 Motor System
(Cerebellum)
• Dysfunction in cerebellum can cause a specific
form of weakness:

• Ataxia: movements lack order because components are not

balanced or coordinated
• Dysmetria: movements overshoot target
• Action tremor: oscillation around motor target; repetitive
correction of previous error due to dysfunction
Advanced Neuroanatomy:
Speech
Higher Order Functions
and Pain
 Neuroanatomy of Speech
• Speech has several components:
– Motor speech: production of words/sentences
– Sensory speech: understanding of speech

• Several levels at which speech can be affected:

– Word production
– Fluency of speech
– Prosody and cadence

• There are cortical and subcortical and brainstem

areas involved in sensory and motor aspects of
speech
 Speech
• Typically the left hemisphere is
responsible for speech and defines
the dominant hemisphere (note
that handedness is not as good as
speech in defining hemispheric
dominance –
• In both right-handed and left-handed
people, the left hemisphere tends to be
dominant
• In about 15% left handed people
there is bilateral speech
representation and in a smaller
number, there is right hemispheric
dominance
 Speech
• Two most important areas that are responsible for
processing of speech are the Broca’s area and
Wernicke’s area
• The tract that connects these two areas together is
called the Arcuate fasciculus
• It is bidirectional in function, meaning that Broca’s area
can send its signal to Wernicke’s area and vice versa
Speech
 Speech
• There are three systems that interact closely in
language perception and production:
1. Language implementation system:
• This system is comprised of the Broca, Wernicke, Insular
cortex and Basal ganglia
• It analyzes incoming auditory signals so as to activate
conceptual knowledge and also ensures phonemic and
grammatical construction

2. Mediational system:
• This system is made up of a number of separate regions in
temporal, parietal and frontal association cortex
• It act as as an intermediary between the implementation
system and conceptual system
 Speech
3. Conceptual system:
• This system is made up of regions in the higher‐order
association cortices, which support conceptual knowledge

• Our current understanding of speech is

contributed by the studying of language
disorders termed “Aphasia”
• There many different types of aphasia: Broca’s,
Wernicke’s, Conduction, Global, Transcortical
Motor, Transcortical Sensory
 Speech (Aphasia)
• Broca's aphasia:
• Broca's aphasia results from damage to Broca's area
(inferior left frontal lobe)
• Patients have non‐fluent speech, it appears laboured
and difficult
• Articulation may be impaired, and the melodic
intonation of normal speech is lacking

• Wernicke's aphasia:
• Wernicke's aphasia is usually caused by damage to the
posterior sector of the left auditory association cortex
• Patients have impaired comprehension and the content
of their speech is fluent, but out of context with the
information
 Speech (Aphasia)
• Conduction aphasia:
• Conduction aphasia is caused by damage to the left
superior temporal gyrus and the inferior parietal lobe
• Patients comprehend simple sentences and produce
intelligible speech but they have difficulties repeat
sentences

• Global aphasia:
• Global aphasia is a combination of Broca, Wernicke and
Conduction aphasia
• Patients with global aphasia have completely lost their
ability to comprehend language, formulate speech, and
repeat sentences
 Speech (Summary of aphasia)
Type of Speech Comprehension Capacity for Other signs Region
aphasia repetition affected
Broca’s Nonfluent, Largely preserved Impaired Right hemiparesis; Left posterior
effortful for single words patient aware of frontal cortex, and
and grammatically defect and may be underlying
simple sentences depressed structures

Wernicke’s Fluent, abundant, Impaired Impaired No motor signs; Left posterior,

well articulated, patient may be superior, and
melodic anxious, agitated, middle temporal
euphoric, or cortex
paranoid

Conduction Fluent with some Intact or largely Impaired Often none; patient Left superior
articulatory defects preserved may have cortical temporal and
sensory loss or supramarginal gyri
weakness in right
arm

Global Scant, nonfluent Impaired Impaired Right hemiplegia Massive left

perisylvian lesion

Transcortical Nonfluent, Intact or largely Intact or largely Sometimes Anterior or superior

explosive preserved preserved rightsided weakness to Broca’s area
Motor

Transcortical Fluent; scant impaired Intact or largely No motor Posterior or inferior

preserved signs to Wernickes’s area
Sensory
37
 Cranial Nerves Involved in Speech
• Cranial nerves which affects speech include:
– Hypoglossal nerve (CN XII): most important cranial nerve because it
controls movement of the tongue
– To a much lesser degree: CN VII, V (if severe motor weakness)

• CN Xll is classified under general somatic efferent (GSM), it

innervates the somatic skeletal muscles of the tongue

• The paralysis of the genioglossus muscle (muscle that runs

from the chin to the tongue) will affect the protrusion of the
tongue which eventually hinders our ability of speech
 Speech (CN)
• CN lX (glossopharyngeal)
• CN IX has five axonal components:
1. SVM (special visceral motor)
2. GVM (general visceral motor)
3. GVA (general visceral afferent)
4. SVA (special visceral afferent)
5. GSA (general somatic afferent)
• Among these components, GVM is the one
that innervates the pharyngeal constrictor
muscles
 Speech (Areas Beyond the Classical Views)

• Damage to left temporal cortex can causes pure naming

defects, while other functions such as grammatical and
phonemic are intact
• Another area is the Insula, this area is important for planning
or coordinating the articulatory movements necessary for
speech
• This area is also damaged in patients with true Broca aphasia
• The Supplementary motor area and the anterior cingulate
region may also play an important role in the initiation and
maintenance of speech
• There are changes in speech noted after cerebellar
injuries: Speech can become ataxic and non‐fluent
 Effect of the Non‐dominant
Hemisphere on Speech
• Non‐dominant hemisphere can have an effect
on speech as well:
– In cases where there is bilateral speech representation
– In cases of true non‐dominance, by affecting the
prosody of speech
– Prosody: Understanding of the emotions in a word
– The functional localization is similar to that seen in the
dominant hemisphere (but localized to the
non‐dominant hemisphere)
 Aprosodia
• Lack of prosody of speech
• Different types:
– Motor
– Sensory
– Global
– Expressive: Changes in pitch and musicality
of tone of speech
 Mutism
• Injury to speech areas does not result in mutism
• Why are some people mute?
• Some syndromes are associated with mutism
– Akinetic mutism – Hypothalamus/third ventricular area injury
– Cerebellar mutism – Seen sometimes in children after cerebellar
surgery
– Bilateral frontal lobe injuries
– Decreased level of consciousness
• Common disorders of speech do not involve mutism but
result in changes in comprehension or motor output of
speech
Higher Cognitive Function
 General Introduction
• Higher cognitive function involves integration of multiple
different motor and sensory processes by multiple brain
regions
• It implies assessment of sensory and motor information
and relation to a context, that involves one or more of
the following:
• Memory
• Emotions
• Pain
• Example:
• What is the difference when you run because there is
a fire behind you or running in a competition?
 Higher Cognitive Functions
– Memory and the brain
– How emotions are processed
– Neuroanatomical areas important in pain
processing
 Memory
• Memory can be classified into two categories:
1. Implicit
2. Explicit
• Lesions of the limbic association areas of the temporal
lobe led to the discovery of the distinction between
implicit and explicit memory
• Implicit memory (Nondeclarative memory)
• Is a type of memory that is recalled unconsciously
• Implicit memory is typically involved in training reflexive
motor or perceptual skills
Memory
• Explicit memory (Declarative
memory)
• Is a type of memory that can
be recalled through
deliberate and conscious
effort
• It is also associated with
factual knowledge of people,
places and things and what
these fact means
• Explicit memory is highly
flexible
 Memory
• Explicit memory can be further classified into:
1. Semantic (memory for facts)
2. Episodic (memory for events and personal experience)

• Knowledge stored as explicit memory is first acquired

through processing in one or more of the three polymodal
association cortices (the prefrontal, limbic, and
parieto‐occipital‐ temporal cortices) that synthesize visual,
auditory and somatic information
• From there the information is conveyed in series to the
parahippocampal and perirhinal cortices, then the entorhinal cortex,
the dentate gyrus, the hippocampus, the subiculum and finally back to
the entorhinal cortex
• From the entorhinal cortex the information is sent back to the
parahippocampal and perirhinal corteces and finally back to the
polymodal association areas of the neocortex
 Memory (Explicit)
• There are three characteristics about explicit
memory:
– There is no single region for memory storage
– Any item of knowledge has multiple representations in
the brain, each of which corresponds to a different
meaning and can be accessed independently
– Both semantic and episodic knowledge are the results
of 4 related but distinct processes: encoding,
consolidation, storage, and retrieval
 Memory (Implicit)
Implicit memory can be either associative or non‐associative:
• Associative
• Requires subjects to learn about the relationship between two stimuli or
between a behaviour and a stimulus
• There are two forms of associative learning: classical conditioning and
operant conditioning
• Non‐associative
• Learning results when an animal or a person is exposed once or
repeatedly to a single type of stimulus
• There are two forms of non‐associative learning: habituation and
sensitization
 Pain
• Pain is a complex perception that we perceive when the
noxious stimulus is present and is influenced by prior
experience
• It involves the following:
• Perception of pain (sensory)
• Reaction to pain (motor)
• Emotional reaction to pain (eg. crying)
• Autonomic reaction
• Changes in mood (sadness, altered awareness of surroundings, difficulty in
processing of new information, particularly in chronic pain syndromes)
• Typically we describe pain in the following manner:
• Nociceptive pain
• Neuropathic pain
• Acute vs chronic pain
 Pain
• Neurons in several regions of the cerebral cortex
respond selectively to nociceptive input
• Two important areas involved in pain
processing:
1. The cingulate gyrus is responsible for processing the
emotional component of pain
2. The insula receives direct projections from the medial
thalamic nuclei and from the ventral and posterior medial
thalamic nuclei
 Insula
• The insular cortex processes information on the
internal state of the body, contributing to the
autonomic component of the overall pain response
• Patients with asymbolia for pain, which is a syndrome
that results in the lesions of the insular cortex, perceive
noxious stimuli as painful and can distinguish sharp
from dull pain
• However, these patients don’t seem to be able to display
proper emotional responses when pain is perceived
• Thus, this suggests that the insular cortex is responsible for
integrating the sensory, affective, and cognitive components
of pain
 The Cingulate Gyrus
• Large C‐shaped gray matter in the medial aspect of
the hemispheres
• It lies dorsal and superior to the corpus callosum
• Involved in a number of higher functions including:
– Pain
– Emotions
– Sadness
• Classified into the following areas:
– Anterior cingulate cortex (ACC): Broadmann areas 25, 24, 32
– Middle cingulate cortex (MCC): Areas 23, 31
– Posterior cingulate cortex (PCC): Area 23
– Retrosplenial cingulate: Areas 29, 30
 The Cingulate Gyrus
– Anterior cingulate area (ACC): Broadmann areas 25, 24,
32
• Includes subgenual and pregenual cingulate areas
• The dorsal ACC is connected with the prefrontal cortex, the
parietal cortex and the motor system
• The ventral ACC is connected with the amygdala, the
hypothalamus and the anterior insula
– Subgenual ACC – thought to be involved in autonomic control
– Pregenual ACC – emotion/autonomic integration
• ACC has also an important role in neuropsychiatric diseases,
including stress and panic disorders, depression and
obsessive‐compulsive disorders
 The Cingulate Gyrus
– Middle cingulate cortex (MCC):
• Anterior MCC
• Posterior MCC
– Each has a different involvement in the processing of emotions
– aMCC – active during fear
– pMCC – does not appear to respond during simple emotions
• Both regions are important in pain: MCC has an important role in the anticipation of
painful events and processing of acute pain
• Neuropathic pain and chronic pain syndromes can cause decreased activity in MCC
– Posterior cingulate cortex (PCC): Dorsal and ventral areas:
• Predominant role in sensory evaluation: self‐relevance of objects and places
• Receives input from the visual areas
– Retrosplenial cingulate:
• Small area, difficult to study; heavily connected with PCC
• Important role in memory; injury to this area can result in profound amnesia
The Cingulate Gyrus
 Sensory System
(Perception of Pain)
• Nociceptive afferent fibers terminate on neurons
in the dorsal horn of the spinal cord
• From the dorsal horn, information is transmitted
to the thalamus and cerebral cortex via five
ascending pathways:
1. Spinothalamic tract
2. Spinoreticular tract
3. Spinomescencephalic tract
4. Cervicothalamic tract
5. Spinohypothalamic tract
• There are two major thalamic nuclei that process
the nociceptive information: lateral and medial
nuclei
 Is the Cortical Sensory System
Responsible for Pain?
• Very complex question
• The sensory system is responsible for the
interpretation of sensation
• However, it appears that the sensory system is not
responsible for the direct perception of pain:
– Injuries in the parietal area do not result in
decreased perception of pain
– Modulation of the parietal area does not
change perception of pain
– Other areas such as the cingulate cortex and
periventricular gray have a much more important role
 Summary
• Overall, motor movement involves integration of the
following systems:
• Supratentorial:
– Cerebral cortex: primary motor, premotor, SMA
– Basal ganglia
– Thalamus
• Infratentorial:
– Brainstem (red nucleus)
– Cerebellum
• Spinal cord
• Peripheral nerves and muscle

• Think therefore of all the different steps required,

between the time you decide to reach for a pen, and
when you carry out the action!