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Learning Objectives
1. Describe the structures involved in psychiatric andneurobehavioral disorders
2. Describe and diagram the basic morphology of thestructures comprising the limbic system
3. Describe and diagram the input-output relationshipsof limbic nuclei
4. Characterize the functions of limbic brain structures
and their underlying mechanisms (where known)5. Develop an understanding of the structural and
functional bases for clinical and behavioral disordersassociated with dysfunctions of the limbic system
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OVERVIEW1. Anatomy of neurobehavior system
1. Overview of the human nervous system2. Anatomy of the brain
1. Cortex cerebri
2. Anatomy of the Limbic system
2. Physiology of neurobehavior system
1. Overview of the motor system1. Pyramidal system
2. Extrapyramidal system
2. Overview of the sensory system
3. Higher functions of the brain1. Intellectual functions of the brain
2. Learning
3. Memory
4. Emotion
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Structures Involved in Psychiatric and
Neurobehavioral Disorders1. Hippocampal formation2. Amygdala
3. Orbitofrontal cortex
4. Cingulate gyrus
5. Hypothalamus
6. Mammilary bodies7. Anterior thalamic nucleus
8. Medial dorsal thalamis nucleus
9. Ventral striatum
10. Frontal lobe
11. Rhinencephalon12. Mesencephalon1. Substantia Nigra
2. Ventral tegmental area (VTA)
3. Formatio reticularis
Limbic system
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OVERVIEW OF THE
FUNCTIONAL ANATOMY
OF THE BRAIN
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Areas of the human cerebral cortex defined by Brodmann in his 1909 publication
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Spatial relationships between basal ganglia, thalamus, and
internal capsule as viewed from the left side.
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OVERVIEW OF THE
LIMBIC SYSTEM
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LIMBIC SYSTEM Is a system that concerns with specific motivated or goal-oriented
behaviors, directly aimed at the maintenance of homeostasis and at the
survival of the individual and of the species (Nieuwenhuys, 1996)
The limbic system receives input from many parts of the cortex and
contains multimodal association areas where various aspects of sensory
experience come together to form a single experience.
The hippocampus, within the limbic system, plays crucial roles in spatial
problem solving and in memory.
Functions:
Maintenance of homeostasis
Motivated and goal-oriented behaviors
Survival of the individual Survival of the species
Learning and memory
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McLeans schema of the evolutionary development of a three-layered
triune brain. Note the location of the limbic system in the middle tier
Brain Circuitry and Signaling in Psychiatry
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Limbic system
Hippocampal
formation
(archicortex,
three layers)
Limbic lobe,
(mesocortex, three
to five layers)
Neocortex,
(five to six layers)
Hippocampus
Dentate gyrus
Amygdala
Phylogenetically
oldest
Newest
Parahippocampal gyrus
Cingulate gyrus
Subcallosal gyrus
Primary motor cortex
Primary sensory cortexAssociation cortex
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Diagram of the structure of the cerebral cortex. A:Golgi neuronal stain. B:Nissl
cellular stain. C:Weigart myelin stain. D:Neuronal connections. Roman and Arabic
numerals indicate the layers of the isocortex (neocortex); 4, external line of
Baillarger (line of Gennari in the occipital lobe); 5b, internal line of Baillarger.
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Stuctures of the Limbic System
1. Hypothalamus
2. Amygdala
3. Septal area4. Hippocampal formation
5. Cingulate gyrus
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Fundamentals of Human Neuropsychology.
A. This medial view of the right hemisphere illustrates the principal structures of the
limbic system, including the cingulate cortex, the hippocampus, and theamygdala.
B. A model of the human limbic system and its major structures. Note: As proposed
by Papez, the limbic system forms a circuit in which the hypothalamus
(mammillary bodies) connect to the hippocampus through the cingulate gyrus,
and the hippocampus connects to the hypothalamus through the fornix. (After
Hamilton, 1976.)
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Structure Connections
Dentate gyrusFrom entorhinal cortex (via perforant pathway and alvear pathway)
To hippocampus (via mossy fibers)
Hippocampus From dentate gyrus (via mossy fibers), septum (via fornix), limbic lobe
(via cingulum)
To mamillary bodies, anterior thalamus, septal area, and tuber
cinereum (via fornix); subcallosal area (via longitudinal striae)Septal area From olfactory bulb, amygdala, fornix
To medial forebrain bundle, hypothalamus, habenula
Amygdala From primitive temporal cortex and sensory association cortex, opposite
amygdala (via anterior commissure)
To hypothalamus (direct amygdalofugal pathway), septal area, andhypothalamus (via stria terminalis)
Some Limbic System Connections
Clinical Neuroanatomy, Waxman,25thed.
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Medial aspect of the right hemispherium showing the corpus callosum.
CCg = Genu; CCb = Body; CCs = Splenium
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Schematic drawing of the major anatomical structures of the limbic system.
Note: The cingulated and parahippocampal gyri form the limbic lobe, a rim
of tissue located along the junction of the diencephalons and the cerebral
hemispheres. n, nucleus.
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Upper cortex and white matter tracts of the brain removed, revealing
the close relationship of the limbic system (hippocampus and fornix)
and striatum in the center of the brain.
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Schema depicting dorsal view of
connections of the amygdala:
14 = olfactory structures,
5 = anterior commissure,
6 = olfactory tubercle,7 = limen insulae,
8 = diagonal band of Broca,
9 = inferior thalamic peduncle,
10 = medial telencephalic
fasciculus,
11 = ventral amygdalofugal pathway,
1217 = amygdaloid nuclei,
18 = lateral hypothalamic area,
1920 = nucleus and stria
medullaris,
21 = stria terminalis,
22 = habenular commissure,
23 = septal nuclei.
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Clinical Neuroanatomy, Waxman,25thed.
Diagram of the principal connections of the limbic system. Olfactory and
amygdaloid connections.
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Clinical Neuroanatomy, Waxman,25thed.
Diagram of the principal connections of the limbic system. Hippocampal
system and great limbic lobe.
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Schematic llustration of the location of the limbic system between
the diencephalon and the neocortexClinical Neuroanatomy, Waxman,25thed.
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Clinical Neuroanatomy, Waxman,25thed.
This limbic lobe consists of a ring of cortex outside the corpus
callosum, largely made up of the subcallosal and cingulate gyri
as well as the parahippocampal gyrus.
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Schematic illustration (left oblique view) of the position of
hippocampal formation in the left hemisphere
Clinical Neuroanatomy, Waxman,25thed.
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Le Grande Lobe Limbique as adapted from Brocasoriginal 1878 drawing of an
otters brain. Brocascallosal gyrus is now termed the cingulate gyrus.
Ref: Clinical Neuroanatomy.pdf
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HIPPOCAMPAL FIBERS project to the MAMMILLARY BODIES, which, in turn, project
through the MAMMILLOTHALAMIC TRACT to the ANTERIOR NUCLEUS. The anterior
thalamic nucleus then projects to the CINGULATE GYRUS, and the axons of the cingulate
gyrus then project back to the HIPPOCAMPAL FORMATION.
Papez circuit
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Ref: Clinical Neuroanatomy.pdf
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Ref: Clinical Neuroanatomy.pdf
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RHINENCEPHALON AND
OLFACTORY SYSTEM
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Olfactory System
Olfaction (the sense of smell) is
one of the oldest senses from a
phylogenetic point of view.
The olfactory system constitutes
an important input to the limbic
system.
The olfactory receptorsare
specialized neurons located in the
olfactory mucous membrane,a
portion of the nasal mucosa.
The axons of the olfactory
receptors travel to the olfactory
bulb.
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The olfactory nerve (lateral view)
Clinical Neuroanatomy, Waxman,25thed.
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Olfactory System,cont.
Within the olfactory bulb,
the olfactory receptor axons
terminate in specialized
synaptic arrangements
(termed glomeruli) on the
dendrites of mitral cells.
Olfactory neurons
expressing a specific
odorant receptor (and thus
responsive to a specific
odorant stimulus) projectprecisely to a small number
of glomeruli within the
olfactory bulb.
Olf S
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Olfactory System,cont
The mitral cells of the olfactory bulb send their
axons posteriorly via the olfactory tracts(also
termed the medialand lateral olfactory stria) to
the olfactory projection area in the cortex. The lateral olfactory striais the projection bundle
of fibers that passes laterally along the floor of the
lateral fissure and enters the olfactory projection
areanear the uncus in the temporal lobe (the
pyriform, entorhinalcortex and parts of the
amygdala.)The pyriform cortex projects, in turn,
via the thalamus to the frontal lobe, where
conscious discrimination of odors presumably
occurs.
The small medial olfactory striapasses medially
and up toward the subcallosal gyrus (the anterior
olfactory nucleus)which sends its axons back to
the olfactory bulbs on both sides, presumably aspart of a feedback circuit that modulates the
sensitivity of olfactory sensation.
Other olfactory fibers reach the anterior
perforated substance to serve olfactory reflex
reactions.
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1. Anosmia (loss of smell):
1. Nasal infection
2. Head trauma that damages the cribrous plate
2. Olfactory (or uncinate) hallucination:
1. Might be a sign of temporal lobe tumor
Olfactory System,Dysfunction
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HIPPOCAMPAL
FORMATION
Structures of the Hippocampal Formation
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Structures of the Hippocampal Formation
1. Hippocampus proper(also called Ammon's horn) :
extends the length of the floor of the
inferior horn of the lateral ventricle
continuous with the fornix below the
splenium of the corpus callosum.
2. Dentate gyrus : is a thin, scalloped strip of cortex that lies
on the upper surface of the
parahippocampal gyrus.
serves as an input station for the
hippocampal formation.
receives inputs from many cortical regions
that are relayed to it via the entorhinal
cortex.
The cells of the dentate gyrus project to the
hippocampus.
3. Subiculum
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Schematic illustration of the major connections to, within, and from the
hippocampal formation. Dentate granule cells (DG) project to pyramidal neurons
in the hippocampus. CA1through CA4are sectors of the hippocampus
Clinical Neuroanatomy, Waxman,25thed.
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Clinical Neuroanatomy, Waxman,25thed.
Schematic illustration of pathways between the hippocampal formation and
the diencephalon. Notice the presence of a loop (Papez circuit), including the
parahippocampal gyrus, hippocampus, mamillary bodies, anterior thalamus,
and cingulate gyrus. Notice also that the neocortex feeds into this loop
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Hippocampal formation in
relation to other limbic
structures.
A, amygdala;
AC, anterior commissure;AN, anterior nucleus of the
thalamus;
B-F, basofrontal region;
CC, corpus callosum (b,
body; g, genu; s, splenium);
CG, cingulate gyrus;E-RC, entorhinal cortex;
F, fornix;
Fm, fimbria;
HF, hippocampal formation;
IG, indusium griseum;
MB, mammillary bodies;MTT, mammillothalamic
tract;
S, septal area;
T, thalamus.
Fib th t f th FORNIX
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Fibers that form the FORNIX
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Diagram illustrates the histological appearance of the cell layers within the
hippocampus and loci of the hippocampal fields, dentate gyrus, and subicular cortex.
CA1-CA4 denote the four sectors of the hippocampus
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Semischematic diagram illustrates: (1) inputs from the entorhinal region, which include
the perforant and alvear pathways; (2) internal circuitry, which includes the connections of
the mossy fibers and Schaffer collaterals; and (3) efferent projections of the hippocampal
formation through the fimbria-fornix system of fibers.
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Major projection targets of the hippocampal formation. The primary output is through the fornix to
diencephalon (i.e., medial hypothalamus, mammillary bodies, and anterior thalamic nucleus) via the
postcommissural fornix and to the septal area via the precommissural fornix. Other connections shown
include efferent fibers that synapse in entorhinal cortex, which, in turn, project to amygdala and
cingulate gyrus
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OFC, orbitofrontal cortex
FAC, Frontal association cortex
PMC, premotor cortex
AAC,auditory association cortex
SAC,somatosensory association cortex
SPL , superior parietal lobuleIPL, inferior parietal lobule
TAC, temporal association cortex,
VAC, visual association cortex
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BFC, basal frontal cortex
OFC, orbitofrontal cortex
FAC, Frontal association cortex
PMC, premotor cortex
CG, cingulate gyrus
CC, corpus callosum
PAC, parietal association cortex
SAC,somatosensory association cortexTAC, temporal association cortex,
VAC, visual association cortex
A, amygdala
H, hippocampus
E, entorhinal cortex
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HIPPOCAMPAL AFFERENTS HIPPOCAMPAL EFFERENTS
LgF, longitudinal fissure
PCS, precentral sulcus
CS, central sulcus
LF, lateral fissure
STS, superior temporal sulcus
MTS, middle temporal sulcus
ITS, inferior temporal sulcus
CoS, collateral sulcus
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HYPOTHALAMUS
A. The approximate boundaries of the
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pp
anterior, middle, and posterior
divisions of the Hypothalamus
B. The medial and lateral zones of the
hypothalamus(shaded). Hypothalamic
cells adjacent to the third ventricle is
paraventricular zone.
Abbreviations:A, amygdala;
AC, anterior commissure;
AcN, accumbens nucleus;
CN, caudate nucleus;
CP, cerebral peduncles;Fc, columns of the fornix;
Fcrus, crus of fornix;
Inf, infundibulum;
MB, mammillary body;
OC, optic chiasm;
ON, optic nerve;
OT, optic tract;
P, putamen;
Pit, pituitary gland;
S, septal nuclei;
SN, substantia nigra;
SubT, subthalamus;
T, thalamus.
HYPOTHALAMIC
PITUITARY CONNECTIONS.
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The posterior portion of the pituitary
(neurohypophysis) is innervated by
hypothalamic neurons that transport the
hypothalamic hormones (oxytocin and
vasopressin) down their axons to bereleased into capillary beds of the
posterior pituitary from where they enter
the general circulation. By contrast, the
capillary beds of the anterior pituitary
(adenohypophysis) are supplied with
hypothalamic hormones (eitherreleasing or inhibitory factors) via a
blood portal system from capillary beds in
the hypothalamus itself. Once released
into the adenohypophysis, these
hypothalamic hormones then stimulate
pituitary cells to synthesize and secrete
their own (pituitary) hormones, which
then are released into the bloodstream.
Note: Some hypothalamic hormones
inhibit the production/secretion of
pituitary hormones.
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SEPTAL AREA
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Topographically organized projections from the hippocampal formation to the
septal area (left side) and topographically arranged efferent projections from
the diagonal band of Broca to the hippocampal formation (right side).
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AMYGDALA
AMYGDALA
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AMYGDALA
The amygdala (amygdaloid
nuclear complex) is a gray
matter mass that lies in themedial temporal pole
between the uncus and the
parahippocampal gyrus.
It is situated just anterior tothe tip of the anterior horn of
the lateral ventricle.
Its fiber connections include :
1. the semicircular stria terminalistothe septal area, preoptic areas and
anterior hypothalamus.
2. amygdalofugal pathwayto the
middle portion of the
hypothalamus.
Nuclei of Amygdala
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Nuclei of Amygdala Two distinct groups of neurons:
1. the large basolateral nuclear group.
receives higher-order sensory information from association areas
in the frontal, temporal, and insular cortex.
Axons run back from the amygdala to the association regions of
the cortex.
also connected, via the stria terminalis and the amygdalofugalpathway, to the ventral striatum and the thalamus.
2. the smaller corticomedial nuclear group.
The corticomedial nuclear group of the amygdala, located close to
the olfactory cortex, is interconnected with it as well as the olfactory
bulb.
Connections also run, via the stria terminalis and amygdalofugal
pathway, to and from the brain stem and hypothalamus.
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the organization of the nuclei of the amygdala
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The major efferent projections of the amygdala. One principal output includes the stria terminalis,
which projects to the bed nucleus of the stria terminalis and to the rostro-caudal extent of the
medial hypothalamus. Fibers from the bed nucleus also supply similar regions of the
hypothalamus. Another important output to the hypothalamus and midbrain PAG uses the ventral
amygdalofugal pathway. Other fibers pass rostrally from the amygdala to the prefrontal cortex.
Dysfunction of the Limbic System
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Stimulation alters somatic motor responses, leading
to bizarre eating and drinking habits, changes insexual and grooming behavior, and defensive
postures of attack and rage.
There can be changes in autonomic responses,
altering cardiovascular or gastrointestinal function,
and in personality, with shifts from passive to
aggressive behavior.
Damage to some areas of the limbic system may alsoprofoundly affect memory.
Dysfunction of the Limbic System
Klver-Bucy Syndrome
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y y
Occurs in patients with bilateral temporal lobe
lesions. The major characteristics of this syndrome are
hyperorality(a tendency to explore objects by placing
them in the mouth together with the indiscriminate eating
or chewing of objects and all kinds of food); hypersexuality,sometimes described as a lack of sexual
inhibition;
psychic blindness,or visual agnosia, in which objects are
no longer recognized; presumably results from damage tothe amydala.
personality changes,usually with abnormal passivity or
docility.
Temporal Lobe Epilepsy
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Temporal Lobe Epilepsy
The temporal lobe (especially the hippocampus and amygdala) has a lower
threshold for epileptic seizure activity than the other cortical areas.
Seizures that originate in these regions, called psychomotor (complex
partial) seizures,differ from the jacksonian seizures that originate in or
near the motor cortex.
Temporal lobe epilepsy may include abnormal sensations, especially
bizarre olfactory sensations, sometimes called uncinate fits; repeated
involuntary movements such as chewing, swallowing, and lip smacking;
disorders of consciousness; memory loss; hallucinations; and disorders of
recall and recognition.
Etiology:
tumor (eg, astrocytoma or oligodendroglioma) may be responsible,
glial scar formation after trauma to the temporal poles may trigger seizures.
Although anticonvulsant drugs are often given to control the seizures, they
may be ineffective. In these cases, neurosurgical removal of the seizure
focus in the temporal lobe may provide excellent seizure control.
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PREFRONTAL CORTEX
PREFRONTAL CORTEX
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PREFRONTAL CORTEX
Located in front of the motor cortex. This is one of
the anatomic structures that distinguishes humansfrom other mammal.
Comprises 29% of the total cortex.
Dysfunction in the PFC is implicated as a possiblesource of pathology in many psychiatric disorders
depression, schizophrenia, anxiety, and attention
deficit hyperactivity disorder (ADHD) as well as anger
and violence.
FRONTAL CORTEX
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The frontal lobe can be divided into three sections.
1. The first, occupying the precentral gyrus, is the PRIMARY
MOTOR CORTEX. Conscious movements are mediated in theprimary motor cortex.
2. the second, lying just anterior to it, is the PREMOTOR
CORTEX. The planning of complex actions occurs in the
premotor cortex.3. The third area, is the PREFRONTAL CORTEX. The prefrontal
cortex is the portion of the frontal cortex lying anterior to
the premotor and primary motor cortices.
1. the executive prefrontal cortex, occupying the dorsal and lateralaspects of the prefrontal cortex;
2. the paralimbic prefrontal cortex, occupying the orbital and medial
aspects of the prefrontal cortex;
3. the anterior cingulate area.
Executive/Dorsolateral Prefrontal Cortex
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Executive/Dorsolateral Prefrontal Cortex
The dorsolateral prefrontal cortex receives innervation from
both the parietal and the temporal association cortices. The dorsolateral prefrontal cortex is responsible for executive
aspects of cognition.
cognitive speed and flexibility,
task sequencing,
higher-order attention,
working memory.
The dorsolateral prefrontal cortex projects to the premotor
cortex lying just posterior to it, suggesting its role in thetransformation of sensory information into preparation for
movement.
Paralimbic/Orbitofrontal Cortex
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Paralimbic/Orbitofrontal Cortex
Regarded as an extension of the limbic system.
It is involved in complex aspects of human behavior, such as
regulation of IMPULSES, MOOD, AND PERSONALITY.
The orbitofrontal cortex receives innervation from the
mediodorsal nucleus of the thalamus, which in turn receives
input from almost all other cortical structures.
The orbitofrontal cortex has reciprocal connections to the
amygdala both through the ventral amygdalofugal tract and
through the stria terminalis. These pathways provide the
orbitofrontal cortex with access to all information necessary
to respond to the environment with respect to themotivational and cognitive state of the individual.
Anterior Cingulate Gyrus
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Anterior Cingulate Gyrus
The cingulate gyrus spans the medial surface of the brain from
the prefrontal cortex to the parieto-occipital junction.
Providing a bridge between the systems performing
emotional, cognitive, motor, and sensory processing.
The anterior cingulate is involved in attention, drive,
motivation, memory, and initiation of speech.
Dysfunction causes:
Apathy.
Akinetic mutism in which the person is alert but has
tremendous inertia, with little spontaneous movement or
speech.
Obsessive compulsive disorder
Basal Ganglia and Ventral Striatum
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Basal Ganglia and Ventral Striatum
The striatum consists of the caudate and the putamen.
The more dorsal components of the striatum are associated with theextrapyramidal system and motor control.
The more ventral areas are involved in motivation, cognition, and
emotion.
At their ventralmost extent, the caudate and putamen appear to merge
into a single structure, called the nucleus accumbens. The striatum is composed of
Striosomes : acetylcholinesterase-poor patches rich in
opiate receptors.
have high levels of D1 receptors and prominent limbic input.
mostly involved in cognitive and behavioral processing.
An acetylcholinesterase-rich matrix.
rich in D2 receptors with close connections to the substantia nigra.
whereas the matrix is a major component of the extrapyramidal motor
system.
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PICTURES OF THE LIMBIC SYSTEM
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From : Human Hippocampus
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1. hippocampus;
2. parahippocampal gyrus;
3. fusiform gyrus;
4. inferior temporal gyrus;
5. middle temporal gyrus;
6. superior temporal gyrus;
7. lateral fissure;
8. postcentral gyrus;
9. central sulcus;
10. precentral gyrus;
11. superior frontal gyrus;
12. cingulate gyrus;
13. corpus callosum;
14. lateral ventricle;
14. caudate nucleus;15. thalamus;
16. putamen;
17. temporal (inferior) horn
of the lateral ventricle;
18. red nucleus;
19. substantia nigra;
20. pons;
21. tentorium cerebelli;
22. ambient cistern
From : Human Hippocampus
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Intraventricular aspect of
the hippocampus.(The temporal horn has been opened
and the choroid plexuses removed.)
1. hippocampal body2. head and digitationes
hippocampi (internal
digitations)
3. hippocampal tail
4. fimbria
5. crus of fornix
6. Subiculum7. splenium of the corpus
callosum;
8. calcar avis
9. collateral trigone
10. collateral eminence
11. uncal recess of thetemporal horn
From : Human Hippocampus
1. anterior paraolfactory sulcus(subcallosal sulcus)
2. cingulate sulcus
3 b i t l l
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3. subparietal sulcus
4. anterior calcarine sulcus;
5. collateral sulcus
6. rhinal sulcus. Limbic gyrus:
7. subcallosal gyrus;
8. posterior paraolfactory sulcus;
9. cingulate gyrus;
10. isthmus;
11. parahippocampal gyrus,
posterior part; 11,
parahippocampal gyrus, anterior
part (piriform lobe).
12. entorhinal area;
13. ambient gyrus;
14. semilunar gyrus;15. prepiriform cortex. Intralimbic
gyrus:
16. prehippocampal rudiment; 16,
paraterminal gyrus;
17. indusium griseum.
Hippocampus:
18. gyrus dentatus;
19. cornu Ammonis;
20. gyri of Andreas Retzius;21. fimbria (displaced upwards,
arrows);
22. uncal apex;
23. band of Giacomini;
24. uncinate gyrus;
25. anterior perforated substance;
26. anterior commissure;
27. fornix;
28. corpus callosumFrom : Human Hippocampus
1. anterior paraolfactory sulcus(subcallosal sulcus)
2. cingulate sulcus
3 b i t l l
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3. subparietal sulcus
4. anterior calcarine sulcus;
5. collateral sulcus
6. rhinal sulcus. Limbic gyrus:
7. subcallosal gyrus;
8. posterior paraolfactory sulcus;
9. cingulate gyrus;
10. isthmus;
11. parahippocampal gyrus,
posterior part; 11,
parahippocampal gyrus, anterior
part (piriform lobe).
12. entorhinal area;
13. ambient gyrus;
14. semilunar gyrus;15. prepiriform cortex. Intralimbic
gyrus:
16. prehippocampal rudiment; 16,
paraterminal gyrus;
17. indusium griseum.
Hippocampus:
18. gyrus dentatus;
19. cornu Ammonis;
20. gyri of Andreas Retzius;21. fimbria (displaced upwards,
arrows);
22. uncal apex;
23. band of Giacomini;
24. uncinate gyrus;
25. anterior perforated substance;
26. anterior commissure;
27. fornix;
28. corpus callosumFrom : Human Hippocampus
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Schematic view of frontal-subcortical-thalamic circuits, shown at the level of the
pallidum and thalamus. The extrapyramidal motor circuit involves the globus pallidus
interna (GPi) and ventral lateral nucleus of the thalamus (VLo) and connects to the
supplementary motor cortex. The related circuit of the ventral striatum involves the
ventral pallidum (VP) and mediodorsal nucleus of the thalamus (MD) and connects to
the prefrontal cortex. IC = internal capsule; III = third ventricle; R = reticular nucleus of
the thalamus; GPe = globus pallidus externa.
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