The Brain

Muse spring 2430

lecture 15
7/12/10
Embryonic Development

• Neural plate forms from ectoderm

• Neural plate invaginates to form a neural
groove and neural folds
 Surface
Head
ectoderm

Neural
plate

Tail
1 The neural plate forms from surface ectoderm.

Figure 12.1, step 1

(a) (b) Primary brain
Neural vesicles
tube

Anterior Prosencephalon
(rostral) (forebrain)

Mesencephalon
(midbrain)

Rhombencephalon
(hindbrain)

Posterior
(caudal)
Figure 12.2a-b
(c) Secondary brain (d) Adult brain (e) Adult
vesicles structures neural canal
regions

Cerebrum: cerebral Lateral

Telencephalon hemispheres (cortex, ventricles
white matter, basal nuclei)

Diencephalon
Diencephalon (thalamus, hypothalamus, Third ventricle
epithalamus), retina

Cerebral
Mesencephalon Brain stem: midbrain
aqueduct

Metencephalon Brain stem: pons

Cerebellum Fourth
ventricle
Myelencephalon Brain stem: medulla
oblongata
Spinal cord Central canal

Figure 12.2c-e
Anterior (rostral) Posterior (caudal)
Metencephalon
Mesencephalon Midbrain
Diencephalon Flexures
Cervical
Telencephalon
Myelencephalon Spinal cord
(a) Week 5

Figure 12.3a
 Cerebral hemisphere
Outline of diencephalon
Midbrain
Cerebellum
Pons
Medulla oblongata
Spinal cord
(b) Week 13

Figure 12.3b
 Cerebral
hemisphere
Cerebellum
Pons
Medulla
oblongata
(c) Week 26 Spinal cord
Figure 12.3c
 Cerebral
hemisphere

Diencephalon
Cerebellum
Brain stem
• Midbrain
• Pons
• Medulla
(d) Birth oblongata

Figure 12.3d
Ventricles of the Brain

• Contain cerebrospinal fluid

• Two C-shaped lateral ventricles in the cerebral
hemispheres
• Third ventricle in the diencephalon
• Fourth ventricle in the hindbrain, dorsal to the
pons, develops from the lumen of the neural
tube
 Lateral ventricle
Septum pellucidum
Anterior horn Posterior
horn
Inferior
horn

Interventricular
Lateral
foramen
aperture Third ventricle Median
Inferior horn aperture
Cerebral aqueduct Lateral
Fourth ventricle aperture
Central canal
(a) Anterior view (b) Left lateral view

Figure 12.5
Cerebral Hemispheres

• Surface markings
• Ridges (gyri), shallow grooves (sulci), and deep
grooves (fissures)
• Five lobes
• Frontal
• Parietal
• Temporal
• Occipital
• Insula
Cerebral Hemispheres

• Surface markings
• Central sulcus
• Separates the precentral gyrus of the frontal lobe
and the postcentral gyrus of the parietal lobe
• Longitudinal fissure
• Separates the two hemispheres
• Transverse cerebral fissure
• Separates the cerebrum and the cerebellum

PLAY Animation: Rotatable brain

 Precentral Central
gyrus sulcus Postcentral
Frontal gyrus Parietal lobe
lobe
Parieto-occipital sulcus
(on medial surface
of hemisphere)
Lateral sulcus
Occipital lobe
Temporal lobe
Transverse cerebral fissure
Cerebellum
Pons
Fissure Medulla oblongata
(a deep Spinal cord
sulcus)
Gyrus
Cortex (gray matter)
Sulcus
White matter

(a)
Figure 12.6a
 Central
Frontal lobe sulcus

Gyri of insula
Temporal lobe
(pulled down)

(b)
Figure 12.6b
 Anterior

Longitudinal Frontal lobe

fissure

Cerebral veins
and arteries Parietal
covered by lobe
arachnoid
mater
Left cerebral Right cerebral
hemisphere hemisphere
Occipital
lobe
(c) Posterior
Figure 12.6c
 Left cerebral
hemisphere

Transverse
Brain stem cerebral
fissure
Cerebellum
(d)
Figure 12.6d
Cerebral Cortex

• Thin (2–4 mm) superficial layer of gray matter

• 40% of the mass of the brain
• Site of conscious mind: awareness, sensory
perception, voluntary motor initiation,
communication, memory storage, understanding
• Each hemisphere connects to contralateral side of
the body
• There is lateralization of cortical function in the
hemispheres
Functional Areas of the Cerebral Cortex

• The three types of functional areas are:

• Motor areas—control voluntary movement
• Sensory areas—conscious awareness of
sensation
• Association areas—integrate diverse
information
• Conscious behavior involves the entire cortex
Motor Areas

• Primary (somatic) motor cortex

• Premotor cortex
• Broca’s area
• Frontal eye field
Motor areas Central sulcus Sensory areas and related
Primary motor cortex association areas
Premotor cortex Primary somatosensory
cortex Somatic
Frontal eye field Somatosensory sensation
association cortex
Broca’s area
(outlined by dashes) Gustatory cortex
Taste
(in insula)
Prefrontal cortex
Working memory Wernicke’s area
for spatial tasks (outlined by dashes)
Executive area for
task management
Working memory for Primary visual
object-recall tasks cortex
Visual Vision
Solving complex,
multitask problems association
area
Auditory
association area Hearing
Primary
auditory cortex
(a) Lateral view, left cerebral hemisphere

Primary motor cortex Motor association cortex Primary sensory cortex

Sensory association cortex Multimodal association cortex

Figure 12.8a
Primary Motor Cortex

• Large pyramidal cells of the precentral gyri

• Long axons  pyramidal (corticospinal) tracts
• Allows conscious control of precise, skilled,
voluntary movements
• Motor homunculi: upside-down caricatures
representing the motor innervation of body
regions
 Posterior

Motor
Motor map in Anterior
precentral gyrus

Toes

Jaw

Tongue Primary motor

cortex
Swallowing (precentral gyrus)

Figure 12.9
Premotor Cortex

• Anterior to the precentral gyrus

• Controls learned, repetitious, or patterned
motor skills
• Coordinates simultaneous or sequential
actions
• Involved in the planning of movements that
depend on sensory feedback
Broca’s Area

• Anterior to the inferior region of the premotor

area
• Present in one hemisphere (usually the left)
• A motor speech area that directs muscles of
the tongue
• Is active as one prepares to speak
Frontal Eye Field

• Anterior to the premotor cortex and superior to

Broca’s area
• Controls voluntary eye movements
Sensory Areas

• Primary somatosensory • Olfactory cortex

cortex
• Gustatory cortex
• Somatosensory
• Visceral sensory area
association cortex
• Vestibular cortex
• Visual areas
• Auditory areas
Motor areas Central sulcus Sensory areas and related
Primary motor cortex association areas
Premotor cortex Primary somatosensory
cortex Somatic
Frontal eye field Somatosensory sensation
association cortex
Broca’s area
(outlined by dashes) Gustatory cortex
Taste
(in insula)
Prefrontal cortex
Working memory Wernicke’s area
for spatial tasks (outlined by dashes)
Executive area for
task management
Working memory for Primary visual
object-recall tasks cortex
Visual Vision
Solving complex,
multitask problems association
area
Auditory
association area Hearing
Primary
auditory cortex
(a) Lateral view, left cerebral hemisphere

Primary motor cortex Motor association cortex Primary sensory cortex

Sensory association cortex Multimodal association cortex

Figure 12.8a
Primary Somatosensory Cortex

• In the postcentral gyri

• Receives sensory information from the skin,
skeletal muscles, and joints
• Capable of spatial discrimination: identification
of body region being stimulated
 Posterior

Sensory
Anterior Sensory map in
postcentral gyrus

Genitals

Primary somato-
sensory cortex Intra-
(postcentral gyrus) abdominal

Figure 12.9
Somatosensory Association Cortex

• Posterior to the primary somatosensory cortex

• Integrates sensory input from primary
somatosensory cortex
• Determines size, texture, and relationship of
parts of objects being felt
Visual Areas

• Primary visual (striate) cortex

• Extreme posterior tip of the occipital lobe
• Most of it is buried in the calcarine sulcus
• Receives visual information from the retinas
Visual Areas

• Visual association area

• Surrounds the primary visual cortex
• Uses past visual experiences to interpret
visual stimuli (e.g., color, form, and movement)
• Complex processing involves entire posterior
half of the hemispheres
Auditory Areas

• Primary auditory cortex

• Superior margin of the temporal lobes
• Interprets information from inner ear as pitch,
loudness, and location
• Auditory association area
• Located posterior to the primary auditory
cortex
• Stores memories of sounds and permits
perception of sounds
OIfactory Cortex

• Medial aspect of temporal lobes (in piriform

lobes)
• Part of the primitive rhinencephalon, along
with the olfactory bulbs and tracts
• (Remainder of the rhinencephalon in humans
is part of the limbic system)
• Region of conscious awareness of odors
Gustatory Cortex

• In the insula
• Involved in the perception of taste
Visceral Sensory Area

• Posterior to gustatory cortex

• Conscious perception of visceral sensations,
e.g., upset stomach or full bladder
Vestibular Cortex

• Posterior part of the insula and adjacent

parietal cortex
• Responsible for conscious awareness of
balance (position of the head in space)
Motor areas Central sulcus Sensory areas and related
Primary motor cortex association areas
Premotor cortex Primary somatosensory
cortex Somatic
Frontal eye field Somatosensory sensation
association cortex
Broca’s area
(outlined by dashes) Gustatory cortex
Taste
(in insula)
Prefrontal cortex
Working memory Wernicke’s area
for spatial tasks (outlined by dashes)
Executive area for
task management
Working memory for Primary visual
object-recall tasks cortex
Visual Vision
Solving complex,
multitask problems association
area
Auditory
association area Hearing
Primary
auditory cortex
(a) Lateral view, left cerebral hemisphere

Primary motor cortex Motor association cortex Primary sensory cortex

Sensory association cortex Multimodal association cortex

Figure 12.8a
 Cingulate Primary
Premotor cortex gyrus motor cortex Central sulcus
Corpus Primary somatosensory
callosum cortex
Frontal eye field Parietal lobe
Somatosensory
Prefrontal
association cortex
cortex
Parieto-occipital
sulcus

Occipital
Processes emotions
lobe
related to personal
and social interactions Visual
Orbitofrontal association
cortex Olfactory bulb area
Olfactory tract Primary
Fornix visual cortex
Temporal lobe Uncus Calcarine sulcus
Primary Parahippocampal
olfactory cortex gyrus
(b) Parasagittal view, right hemisphere

Primary motor cortex Motor association cortex Primary sensory cortex

Sensory association cortex Multimodal association cortex

Figure 12.8b
Multimodal Association Areas

• Receive inputs from multiple sensory areas

• Send outputs to multiple areas, including the
premotor cortex
• Allow us to give meaning to information
received, store it as memory, compare it to
previous experience, and decide on action to
take
Multimodal Association Areas

• Three parts
• Anterior association area (prefrontal cortex)
• Posterior association area
• Limbic association area
Anterior Association Area (Prefrontal
Cortex)
• Most complicated cortical region
• Involved with intellect, cognition, recall, and
personality
• Contains working memory needed for
judgment, reasoning, persistence, and
conscience
• Development depends on feedback from
social environment
Posterior Association Area

• Large region in temporal, parietal, and

occipital lobes
• Plays a role in recognizing patterns and faces
and localizing us in space
• Involved in understanding written and spoken
language (Wernicke’s area)
Limbic Association Area

• Part of the limbic system

• Provides emotional impact that helps
establish memories
Lateralization of Cortical Function

• Lateralization
• Division of labor between hemispheres
• Cerebral dominance
• Designates the hemisphere dominant for
language (left hemisphere in 90% of people)
Lateralization of Cortical Function

• Left hemisphere
• Controls language, math, and logic
• Right hemisphere
• Insight, visual-spatial skills, intuition, and
artistic skills
• Left and right hemispheres communicate via
fiber tracts in the cerebral white matter
Cerebral White Matter

• Myelinated fibers and their tracts

• Responsible for communication
• Commissures (in corpus callosum)—connect
gray matter of the two hemispheres
• Association fibers—connect different parts of
the same hemisphere
• Projection fibers—(corona radiata) connect the
hemispheres with lower brain or spinal cord
 Commissural
Longitudinal fissure fibers (corpus
Superior
callosum)
Lateral
ventricle
Association
Basal nuclei fibers
• Caudate
• Putamen Corona radiata
• Globus Fornix
pallidus
Internal
Thalamus capsule
Gray matter
Third
White matter
ventricle
Projection
Pons
fibers
Decussation
Medulla oblongata of pyramids
(a)

Figure 12.10a
Basal Nuclei (Ganglia)

• Subcortical nuclei
• Consists of the corpus striatum
• Caudate nucleus
• Lentiform nucleus (putamen + globus pallidus)
• Functionally associated with the subthalamic
nuclei (diencephalon) and the substantia nigra
(midbrain)
Fibers of
corona radiata
Caudate
nucleus Thalamus
Lentiform Tail of
Corpus nucleus caudate
striatum • Putamen nucleus
• Globus pallidus
(deep to putamen)
Projection fibers
run deep to
(a) lentiform nucleus

Figure 12.11a
 Anterior
Cerebral cortex
Cerebral white matter
Corpus callosum
Anterior horn
of lateral ventricle
Caudate nucleus
Putamen
Globus Lentiform
pallidus nucleus
Thalamus
Tail of caudate nucleus
Third ventricle
Inferior horn
(b) of lateral ventricle
Posterior
Figure 12.11b (1 of 2)
Cerebral cortex
Cerebral white matter
Corpus callosum
Anterior horn
of lateral ventricle
Caudate nucleus

Lentiform nucleus

Thalamus

Third ventricle
Inferior horn
of lateral ventricle
(b)

Figure 12.11b (2 of 2)
Functions of Basal Nuclei

• Though somewhat elusive, the following are

thought to be functions of basal nuclei
• Influence muscular control
• Help regulate attention and cognition
• Regulate intensity of slow or stereotyped
movements
• Inhibit antagonistic and unnecessary
movements
Diencephalon

• Three paired structures

• Thalamus
• Hypothalamus
• Epithalamus
• Encloses the third ventricle

PLAY Animation: Rotatable brain (sectioned)

Cerebral hemisphere
Septum pellucidum Corpus callosum
Interthalamic Fornix
adhesion Choroid plexus
(intermediate Thalamus
mass of (encloses third
thalamus) ventricle)
Interven- Posterior commissure
tricular Pineal gland
foramen (part of epithalamus)
Anterior Corpora
commissure quadrigemina Mid-
Hypothalamus Cerebral brain
aqueduct
Optic chiasma
Arbor vitae (of
Pituitary gland cerebellum)
Fourth ventricle
Mammillary body
Pons Choroid plexus
Cerebellum
Medulla oblongata
Spinal cord

Figure 12.12
Thalamus

• 80% of diencephalon
• Superolateral walls of the third ventricle
• Connected by the interthalamic adhesion
(intermediate mass)
• Contains several nuclei, named for their
location
• Nuclei project and receive fibers from the
cerebral cortex
 Dorsal nuclei

Medial Lateral Lateral

dorsal posterior

Pulvinar

Anterior Medial
nuclear geniculate
group body
Reticular Lateral
nucleus geniculate
Ventral body
Ventral Ventral postero-
anterior lateral lateral

Ventral nuclei
(a) The main thalamic nuclei. (The reticular nuclei that “cap” the
thalamus laterally are depicted as curving translucent structures.)
Figure 12.13a
Thalamic Function

• Gateway to the cerebral cortex

• Sorts, edits, and relays information
• Afferent impulses from all senses and all parts of the
body
• Impulses from the hypothalamus for regulation of
emotion and visceral function
• Impulses from the cerebellum and basal nuclei to help
direct the motor cortices
• Mediates sensation, motor activities, cortical arousal,
learning, and memory
Hypothalamus

• Forms the inferolateral walls of the third

ventricle
• Contains many nuclei
• Example: mammillary bodies
• Paired anterior nuclei
• Olfactory relay stations
• Infundibulum—stalk that connects to the
pituitary gland
 Paraventricular
nucleus

Anterior Dorsomedial
Fornix
commissure nucleus
Preoptic
Posterior
nucleus hypothalamic
Anterior nucleus
hypothalamic Lateral
nucleus hypothalamic
Supraoptic area
nucleus Ventromedial
Supra- nucleus
chiasmatic Mammillary
nucleus Arcuate body
Optic nucleus
chiasma Pituitary
Infundibulum
gland
(stalk of the
pituitary gland)
(b) The main hypothalamic nuclei.
Figure 12.13b
Hypothalamic Function

• Autonomic control center for many visceral

functions (e.g., blood pressure, rate and force
of heartbeat, digestive tract motility)
• Center for emotional response: Involved in
perception of pleasure, fear, and rage and in
biological rhythms and drives
Hypothalamic Function

• Regulates body temperature, food intake,

water balance, and thirst
• Regulates sleep and the sleep cycle
• Controls release of hormones by the anterior
pituitary
• Produces posterior pituitary hormones
Epithalamus

• Most dorsal portion of the diencephalon;

forms roof of the third ventricle
• Pineal gland—extends from the posterior
border and secretes melatonin
• Melatonin—helps regulate sleep-wake cycles
Cerebral hemisphere
Septum pellucidum Corpus callosum
Interthalamic Fornix
adhesion Choroid plexus
(intermediate Thalamus
mass of (encloses third
thalamus) ventricle)
Interven- Posterior commissure
tricular Pineal gland
foramen (part of epithalamus)
Anterior Corpora
commissure quadrigemina Mid-
Hypothalamus Cerebral brain
Optic chiasma aqueduct
Arbor vitae (of
Pituitary gland cerebellum)
Fourth ventricle
Mammillary body
Pons Choroid plexus
Cerebellum
Medulla oblongata
Spinal cord

Figure 12.12
Brain Stem

• Three regions
• Midbrain
• Pons
• Medulla oblongata
Brain Stem

• Similar structure to spinal cord but contains

embedded nuclei
• Controls automatic behaviors necessary for
survival
• Contains fiber tracts connecting higher and
lower neural centers
• Associated with 10 of the 12 pairs of cranial
nerves
Frontal lobe
Olfactory bulb
(synapse point of
cranial nerve I)
Optic chiasma
Optic nerve (II)
Optic tract
Mammillary body
Midbrain
Pons
Temporal lobe
Medulla
oblongata
Cerebellum
Spinal cord

Figure 12.14
 View (a)
Optic chiasma
Optic nerve (II)
Diencephalon
Crus cerebri of
• Thalamus cerebral peduncles
• Hypothalamus
(midbrain) Thalamus
Diencephalon
Mammillary body Hypothalamus
Midbrain
Oculomotor nerve (III)
Pons Brainstem
Trochlear nerve (IV) Medulla
oblongata
Trigeminal nerve (V)
Middle cerebellar Pons
peduncle Facial nerve (VII)
Abducens nerve (VI)
Vestibulocochlear Glossopharyngeal nerve (IX)
nerve (VIII) Hypoglossal nerve (XII)
Pyramid
Vagus nerve (X)
Ventral root of first
cervical nerve Accessory nerve (XI)
Decussation of pyramids

Spinal cord

(a) Ventral view

Figure 12.15a
Crus cerebri of
Thalamus
cerebral peduncles
(midbrain)
View (b)
Infundibulum Superior colliculus
Pituitary gland Inferior colliculus
Trochlear nerve (IV)
Trigeminal nerve (V) Superior cerebellar peduncle
Pons Middle cerebellar peduncle
Facial nerve (VII)
Inferior cerebellar peduncle
Abducens nerve (VI)
Vestibulocochlear nerve (VIII)
Glossopharyngeal nerve (IX) Olive
Hypoglossal nerve (XII)
Thalamus
Vagus nerve (X) Hypothalamus
Diencephalon

Midbrain
Accessory nerve (XI)
Pons Brainstem
Medulla
oblongata

(b) Left lateral view

Figure 12.15b
 Thalamus
View (c)

Diencephalon
Midbrain
• Superior Corpora
colliculus quadrigemina
• Inferior of tectum
colliculus
Pineal gland • Trochlear nerve (IV)
• Superior cerebellar peduncle
Pons
• Middle cerebellar peduncle
Medulla oblongata
Anterior wall of • Inferior cerebellar peduncle
fourth ventricle • Facial nerve (VII)
• Vestibulocochlear nerve (VIII)
Choroid plexus • Glossopharyngeal nerve (IX)
(fourth ventricle) • Vagus nerve (X)
Dorsal median sulcus • Accessory nerve (XI)
Thalamus
Dorsal root of Hypothalamus
Diencephalon

first cervical nerve Midbrain

Pons Brainstem
(c) Dorsal view Medulla
oblongata

Figure 12.15c
Midbrain

• Located between the diencephalon and the

pons
• Cerebral peduncles
• Contain pyramidal motor tracts
• Cerebral aqueduct
• Channel between third and fourth ventricles
Midbrain Nuclei

• Nuclei that control cranial nerves III (oculomotor) and

IV (trochlear)
• Corpora quadrigemina—domelike dorsal protrusions
• Superior colliculi—visual reflex centers
• Inferior colliculi—auditory relay centers
• Substantia nigra—functionally linked to basal nuclei
• Red nucleus—relay nuclei for some descending
motor pathways and part of reticular formation
 Dorsal Superior
Tectum
Periaqueductal gray colliculus
matter Cerebral aqueduct
Oculomotor
nucleus (III) Reticular formation
Medial
lemniscus
Red
nucleus
Substantia
nigra
Fibers of
pyramidal tract
Ventral Crus cerebri of
(a) Midbrain cerebral peduncle

Figure 12.16a
Pons

• Forms part of the anterior wall of the fourth ventricle

• Fibers of the pons
• Connect higher brain centers and the spinal cord
• Relay impulses between the motor cortex and the
cerebellum
• Origin of cranial nerves V (trigeminal), VI (abducens),
and VII (facial)
• Some nuclei of the reticular formation
• Nuclei that help maintain normal rhythm of breathing
 Fourth
ventricle
Reticular
Superior cerebellar formation
peduncle
Trigeminal main
sensory nucleus
Trigeminal
motor nucleus
Middle
cerebellar
peduncle
Trigeminal Pontine
nerve (V) nuclei
Fibers of
Medial lemniscus pyramidal
(b) Pons tract
Figure 12.16b
Medulla Oblongata

• Joins spinal cord at foramen magnum

• Forms part of the ventral wall of the fourth
ventricle
• Contains a choroid plexus of the fourth
ventricle
• Pyramids—two ventral longitudinal ridges
formed by pyramidal tracts
• Decussation of the pyramids—crossover of
the corticospinal tracts
Medulla Oblongata

• Inferior olivary nuclei—relay sensory

information from muscles and joints to
cerebellum
• Cranial nerves VIII, X, and XII are associated
with the medulla
• Vestibular nuclear complex—mediates
responses that maintain equilibrium
• Several nuclei (e.g., nucleus cuneatus and
nucleus gracilis) relay sensory information
Medulla Oblongata

• Autonomic reflex centers

• Cardiovascular center
• Cardiac center adjusts force and rate of heart
contraction
• Vasomotor center adjusts blood vessel
diameter for blood pressure regulation
Medulla Oblongata

• Respiratory centers
• Generate respiratory rhythm
• Control rate and depth of breathing, with
pontine centers
Medulla Oblongata

• Additional centers regulate

• Vomiting
• Hiccuping
• Swallowing
• Coughing
• Sneezing
 Fourth ventricle
Solitary
Hypoglossal nucleus (XII) Choroid
nucleus
Dorsal motor nucleus plexus Vestibular nuclear
of vagus (X) complex (VIII)
Inferior cerebellar Cochlear
peduncle nuclei (VIII)
Reticular formation

Lateral
nuclear
group Nucleus
Medial ambiguus
nuclear
group Inferior olivary
Raphe nucleus
nucleus Pyramid
Medial lemniscus
(c) Medulla oblongata

Figure 12.16c
The Cerebellum

• 11% of brain mass

• Dorsal to the pons and medulla
• Subconsciously provides precise timing and
appropriate patterns of skeletal muscle
contraction
Anatomy of the Cerebellum

• Two hemispheres connected by vermis

• Each hemisphere has three lobes
• Anterior, posterior, and flocculonodular
• Folia—transversely oriented gyri
• Arbor vitae—distinctive treelike pattern of the
cerebellar white matter
 Anterior lobe
Cerebellar cortex
Arbor
vitae

Cerebellar
peduncles Posterior
• Superior lobe
• Middle Choroid
• Inferior plexus of
Medulla Flocculonodular fourth
oblongata lobe ventricle
(b)
Figure 12.17b
Anterior
lobe

Posterior
lobe

(d) Vermis

Figure 12.17d
Cerebellar Peduncles

• All fibers in the cerebellum are ipsilateral

• Three paired fiber tracts connect the
cerebellum to the brain stem
• Superior peduncles connect the cerebellum to
the midbrain
• Middle peduncles connect the pons to the
cerebellum
• Inferior peduncles connect the medulla to the
cerebellum
Cerebellar Processing for Motor Activity

• Cerebellum receives impulses from the cerebral

cortex of the intent to initiate voluntary muscle
contraction
• Signals from proprioceptors and visual and
equilibrium pathways continuously “inform” the
cerebellum of the body’s position and momentum
• Cerebellar cortex calculates the best way to
smoothly coordinate a muscle contraction
• A “blueprint” of coordinated movement is sent to the
cerebral motor cortex and to brain stem nuclei
Cognitive Function of the Cerebellum

• Recognizes and predicts sequences of events

during complex movements
• Plays a role in nonmotor functions such as
word association and puzzle solving
Functional Brain Systems

• Networks of neurons that work together and

span wide areas of the brain
• Limbic system
• Reticular formation
Limbic System

• Structures on the medial aspects of cerebral

hemispheres and diencephalon
• Includes parts of the diencephalon and some
cerebral structures that encircle the brain
stem
 Septum pellucidum Fiber tracts
connecting limbic
Diencephalic structures system structures
Corpus callosum
of the limbic system
•Fornix
•Anterior thalamic •Anterior commissure
nuclei (flanking
3rd ventricle) Cerebral struc-
tures of the
•Hypothalamus limbic system
•Mammillary
body •Cingulate gyrus
•Septal nuclei
•Amygdala
•Hippocampus
•Dentate gyrus
•Parahippocampal
gyrus
Olfactory bulb

Figure 12.18
Limbic System

• Emotional or affective brain

• Amygdala—recognizes angry or fearful facial
expressions, assesses danger, and elicits the
fear response
• Cingulate gyrus—plays a role in expressing
emotions via gestures, and resolves mental
conflict
• Puts emotional responses to odors
• Example: skunks smell bad
Limbic System: Emotion and Cognition

• The limbic system interacts with the prefrontal

lobes, therefore:
• We can react emotionally to things we
consciously understand to be happening
• We are consciously aware of emotional
richness in our lives
• Hippocampus and amygdala—play a role in
memory
Reticular Formation

• Three broad columns along the length of the

brain stem
• Raphe nuclei
• Medial (large cell) group of nuclei
• Lateral (small cell) group of nuclei
• Has far-flung axonal connections with
hypothalamus, thalamus, cerebral cortex,
cerebellum, and spinal cord
Reticular Formation: RAS and Motor Function

• RAS (reticular activating system)

• Sends impulses to the cerebral cortex to keep
it conscious and alert
• Filters out repetitive and weak stimuli (~99% of
all stimuli!)
• Severe injury results in permanent
unconsciousness (coma)
Reticular Formation: RAS and Motor Function

• Motor function
• Helps control coarse limb movements
• Reticular autonomic centers regulate visceral
motor functions
• Vasomotor
• Cardiac
• Respiratory centers
 Radiations
to cerebral
cortex

Visual
impulses Auditory
impulses
Reticular formation
Ascending general Descending
sensory tracts motor projections
(touch, pain, temperature) to spinal cord

Figure 12.19
Electroencephalogram (EEG)

• Records electrical activity that accompanies

brain function
• Measures electrical potential differences
between various cortical areas
(a) Scalp electrodes are used to record brain wave
activity (EEG).

Figure 12.20a
Brain Waves

• Patterns of neuronal electrical activity

• Generated by synaptic activity in the cortex
• Each person’s brain waves are unique
• Can be grouped into four classes based on
frequency measured as Hertz (Hz)
Types of Brain Waves

• Alpha waves (8–13 Hz)—regular and rhythmic, low-

amplitude, synchronous waves indicating an “idling”
brain
• Beta waves (14–30 Hz)—rhythmic, less regular
waves occurring when mentally alert
• Theta waves (4–7 Hz)—more irregular; common in
children and uncommon in adults
• Delta waves (4 Hz or less)—high-amplitude waves
seen in deep sleep and when reticular activating
system is damped, or during anesthesia; may
indicate brain damage
 1-second interval

Alpha waves—awake but relaxed

Beta waves—awake, alert

Theta waves—common in children

Delta waves—deep sleep

(b) Brain waves shown in EEGs fall into

four general classes.
Figure 12.20b
Brain Waves: State of the Brain

• Change with age, sensory stimuli, brain

disease, and the chemical state of the body
• EEGs used to diagnose and localize brain
lesions, tumors, infarcts, infections,
abscesses, and epileptic lesions
• A flat EEG (no electrical activity) is clinical
evidence of death
Epilepsy

• A victim of epilepsy may lose consciousness,

fall stiffly, and have uncontrollable jerking
• Epilepsy is not associated with intellectual
impairments
• Epilepsy occurs in 1% of the population
Epileptic Seizures

• Absence seizures, or petit mal

• Mild seizures seen in young children where the
expression goes blank
• Tonic-clonic (grand mal) seizures
• Victim loses consciousness, bones are often
broken due to intense contractions, may
experience loss of bowel and bladder control,
and severe biting of the tongue
Control of Epilepsy

• Anticonvulsive drugs
• Vagus nerve stimulators implanted under the
skin of the chest can keep electrical activity of
the brain from becoming chaotic
Consciousness

• Conscious perception of sensation

• Voluntary initiation and control of movement
• Capabilities associated with higher mental
processing (memory, logic, judgment, etc.)
• Loss of consciousness (e.g., fainting or
syncopy) is a signal that brain function is
impaired
Consciousness

• Clinically defined on a continuum that grades

behavior in response to stimuli
• Alertness
• Drowsiness (lethargy)
• Stupor
• Coma
Sleep

• State of partial unconsciousness from which a

person can be aroused by stimulation
• Two major types of sleep (defined by EEG
patterns)
• Nonrapid eye movement (NREM)
• Rapid eye movement (REM)
Sleep

• First two stages of NREM occur during the

first 30–45 minutes of sleep
• Fourth stage is achieved in about 90 minutes,
and then REM sleep begins abruptly
 Awake

REM: Skeletal
muscles (except
ocular muscles
and diaphragm)
are actively
inhibited; most
dreaming occurs.
NREM stage 1:
Relaxation begins;
EEG shows alpha
waves, arousal is easy.

NREM stage 2: Irregular

EEG with sleep spindles
(short high- amplitude
bursts); arousal is more
difficult.
NREM stage 3: Sleep
deepens; theta and
delta waves appear;
vital signs decline.

NREM stage 4: EEG is

dominated by delta
waves; arousal is difficult;
bed-wetting, night terrors,
and sleepwalking may
occur.
(a) Typical EEG patterns

Figure 12.21a
Sleep Patterns

• Alternating cycles of sleep and wakefulness

reflect a natural circadian (24-hour) rhythm
• RAS activity is inhibited during, but RAS also
mediates, dreaming sleep
• The suprachiasmatic and preoptic nuclei of
the hypothalamus time the sleep cycle
• A typical sleep pattern alternates between
REM and NREM sleep
 Awake
REM
Stage 1
Stage 2
Non
REM Stage 3
Stage 4

Time (hrs)
(b) Typical progression of an adult through one
night’s sleep stages

Figure 12.21b
Importance of Sleep

• Slow-wave sleep (NREM stages 3 and 4) is

presumed to be the restorative stage
• People deprived of REM sleep become moody and
depressed
• REM sleep may be a reverse learning process where
superfluous information is purged from the brain
• Daily sleep requirements decline with age
• Stage 4 sleep declines steadily and may disappear
after age 60
Sleep Disorders

• Narcolepsy
• Lapsing abruptly into sleep from the awake
state
• Insomnia
• Chronic inability to obtain the amount or quality
of sleep needed
• Sleep apnea
• Temporary cessation of breathing during sleep
Language

• Language implementation system

• Basal nuclei
• Broca’s area and Wernicke’s area (in the
association cortex on the left side)
• Analyzes incoming word sounds
• Produces outgoing word sounds and grammatical
structures
• Corresponding areas on the right side are
involved with nonverbal language components
Memory

• Storage and retrieval of information

• Two stages of storage
• Short-term memory (STM, or working memory)
—temporary holding of information; limited to
seven or eight pieces of information
• Long-term memory (LTM) has limitless
capacity
 Outside stimuli

General and special sensory receptors

Afferent inputs

Temporary storage
Data permanently
(buffer) in
lost
cerebral cortex

Data selected
Automatic
for transfer Forget
memory

Short-term
memory (STM) Forget

Data transfer
influenced by:
Retrieval Excitement
Rehearsal
Association of
old and new data

Long-term
memory
(LTM) Data unretrievable

Figure 12.22
Transfer from STM to LTM

• Factors that affect transfer from STM to LTM

• Emotional state—best if alert, motivated,
surprised, and aroused
• Rehearsal—repetition and practice
• Association—tying new information with old
memories
• Automatic memory—subconscious information
stored in LTM
Categories of Memory

• Declarative memory (factual knowledge)

• Explicit information
• Related to our conscious thoughts and our
language ability
• Stored in LTM with context in which it was
learned
Categories of Memory

• Nondeclarative memory
• Less conscious or unconscious
• Acquired through experience and repetition
• Best remembered by doing; hard to unlearn
• Includes procedural (skills) memory, motor
memory, and emotional memory
Brain Structures Involved in Declarative
Memory
• Hippocampus and surrounding temporal lobes
function in consolidation and access to
memory
• ACh from basal forebrain is necessary for
memory formation and retrieval
 Thalamus

Basal forebrain
Touch
Prefrontal cortex
Hearing

Vision Taste Smell

Hippocampus
Sensory
Thalamus
input
(a) Declarative
memory circuits
Association Medial temporal lobe Prefrontal
cortex (hippocampus, etc.) cortex

ACh ACh
Basal
forebrain

Figure 12.23a
Brain Structures Involved in Nondeclarative
Memory
• Procedural memory
• Basal nuclei relay sensory and motor inputs to
the thalamus and premotor cortex
• Dopamine from substantia nigra is necessary
• Motor memory—cerebellum
• Emotional memory—amygdala
Sensory and
motor inputs

Association Basal Thalamus Premotor

cortex nuclei cortex

Dopamine Premotor
Substantia cortex
nigra

Thalamus Basal nuclei

Substantia nigra

(b) Procedural (skills) memory circuits

Figure 12.23b
Molecular Basis of Memory

• During learning:
• Altered mRNA is synthesized and moved to axons and
dendrites
• Dendritic spines change shape
• Extracellular proteins are deposited at synapses
involved in LTM
• Number and size of presynaptic terminals may
increase
• More neurotransmitter is released by presynaptic
neurons
Molecular Basis of Memory

• Increase in synaptic strength (long-term

potentiation, or LTP) is crucial
• Neurotransmitter (glutamate) binds to NMDA
receptors, opening calcium channels in
postsynaptic terminal
Molecular Basis of Memory

• Calcium influx triggers enzymes that modify

proteins of the postsynaptic terminal and
presynaptic terminal (via release of retrograde
messengers)
• Enzymes trigger postsynaptic gene activation
for synthesis of synaptic proteins, in presence
of CREB (cAMP response-element binding
protein) and BDNF (brain-derived
neurotrophic factor)
Meninges

• Three layers
• Dura mater
• Arachnoid mater
• Pia mater
 Skin of scalp
Periosteum
Bone of skull
Periosteal Dura
Meningeal mater
Superior Arachnoid mater
sagittal sinus Pia mater
Subdural Arachnoid villus
space Blood vessel
Subarachnoid Falx cerebri
space (in longitudinal
fissure only)

Figure 12.24
Dura Mater

• Strongest meninx
• Two layers of fibrous connective tissue
(around the brain) separate to form dural
sinuses
Dura Mater

• Dural septa limit excessive movement of the

brain
• Falx cerebri—in the longitudinal fissure;
attached to crista galli
• Falx cerebelli—along the vermis of the
cerebellum
• Tentorium cerebelli—horizontal dural fold over
cerebellum and in the transverse fissure
 Superior
sagittal sinus
Falx cerebri
Straight
sinus
Crista galli
Tentorium
of the
ethmoid cerebelli
bone Falx
Pituitary cerebelli
gland

(a) Dural septa

Figure 12.25a
Arachnoid Mater

• Middle layer with weblike extensions

• Separated from the dura mater by the
subdural space
• Subarachnoid space contains CSF and blood
vessels
• Arachnoid villi protrude into the superior
sagittal sinus and permit CSF reabsorption
 Skin of scalp
Periosteum
Bone of skull
Periosteal Dura
Meningeal mater
Superior Arachnoid mater
sagittal sinus Pia mater
Subdural Arachnoid villus
space Blood vessel
Subarachnoid Falx cerebri
space (in longitudinal
fissure only)

Figure 12.24
Pia Mater

• Layer of delicate vascularized connective

tissue that clings tightly to the brain
Superior
sagittal sinus 4

Choroid
plexus Arachnoid villus
Interventricular
foramen Subarachnoid space
Arachnoid mater
Meningeal dura mater
Periosteal dura mater
1
Right lateral ventricle
(deep to cut)
Choroid plexus
3
of fourth ventricle
Third ventricle
1 CSF is produced by the
Cerebral aqueduct choroid plexus of each
Lateral aperture ventricle.
Fourth ventricle 2 CSF flows through the
2 ventricles and into the
Median aperture subarachnoid space via the
Central canal median and lateral apertures.
of spinal cord Some CSF flows through the
central canal of the spinal cord.
3 CSF flows through the
(a) CSF circulation
subarachnoid space.
4 CSF is absorbed into the dural venous
sinuses via the arachnoid villi.

Figure 12.26a
Choroid Plexuses

• Produce CSF at a constant rate

• Hang from the roof of each ventricle
• Clusters of capillaries enclosed by pia mater
and a layer of ependymal cells
• Ependymal cells use ion pumps to control the
composition of the CSF and help cleanse CSF
by removing wastes
Ependymal
cells
Capillary
Section
of choroid
Connective
plexus
tissue of
pia mater

Wastes and CSF forms as a filtrate

Cavity of
unnecessary containing glucose, oxygen,
solutes absorbed ventricle vitamins, and ions
(Na+, Cl–, Mg2+, etc.)
(b) CSF formation by choroid plexuses

Figure 12.26b
Blood-Brain Barrier

• Composition
• Continuous endothelium of capillary walls
• Basal lamina
• Feet of astrocytes
• Provide signal to endothelium for the
formation of tight junctions
 Capillary

Neuron

Astrocyte

a) Astrocytes are the most abundant

CNS neuroglia.

Figure 11.3a