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Limbic System

Affective
Behavior

Emotion
Joy
and
Despair
Introduction

Concept of Limbic System

Broca (1877) - ‘La Grand Lobe Limbique’

Papez (1937) - ‘Limbic Circuit’ - emotion

MacLean (1952) - ‘Limbic System’ - visceral brain

Nauta (1972) - ‘Septo-hypothalamo-mesencephalic continuum’


PAUL BROCA JAMES PAPEZ
(1824-1880) (1883-1958)
Limbic System
- term of Paul MacLean (1952)
- Visceral Brain
- triune concept of brain evolution

Hypothalamus
Nucleus accumbens
amygdaloid nuclear complex
orbitofrontal cortex

Some psychiatric implications on


physiological studies on fronto-
temporal portions of limbic system
(visceral brain). Electroencephalogr
Clin Neurophysiol 4: 407-418, 1952 Paul D. MacLean (1913- )
Main Components of Limbic System

Olfactory
Olfactory
Cortex
Cortex
Septo-(Preoptico)-Hypothalamo-
Septo-(Preoptico)-Hypothalamo-Mesencephalic
MesencephalicContinuum
Continuum
Spinal
Limbic
Limbic Cord
Hippocampal
Hippocampal Septal
Septal Hypothalamus
Hypothalamus Midbrain
Midbrain &
&
Formation
Formation Region
Region Area
Area Brain
Brain
Stem

Amygdaloid
Amygdaloid
Nuclear
Nuclear
Complex
Complex
Limbic System
Expression
of Emotion

Septo-
hypothalamo-
mesencephalic
continuum
Components of the Limbic System (I)

TELENCEPHALON Basal Forebrain Area


Limbic Lobe Substantia Innominata
Gyrus Fornicatus Basal Nucleus of Meynert
Cingulate Gyrus Ventral Pallidum
Parahippocampal Gyrus Nucleus Accumbens (Septi)
Hippocampal Formation Septal Region
Hippocampus Septal Area
Dentate Gyrus Paraterminal Gyrus
Subiculum Subcallosal Gyrus
Hippocampal Rudiments Septal Nuclei
Fasciolar Gyrus Dorsal, Lateral and Medial
Indusium Griseum Septal Nuclei
Olfactory Cortex (Rhinencephalon) Nucleus of Diagonal Band of Broca
- Piriform Lobe Bed Nucleus of Stria Terminalis
Amygdaloid Nuclear Complex
Prepiriform Cortex
Corticomedial Nuclear Group
Periamygdaloid Cortex
Basolateral Nuclear Group
Entorhinal Cortex Central Nucleus
Components of the Limbic System (II)

DIENCEPHALON
Thalamus, Limbic Mammillary Region
Anterior Nuclear Group Lateral, Medial and Intermediate
Mediodorsal (MD) Nucleus Mammillary Nucleus
Midline Nuclei Posterior Nucleus
Hypothalamus Epithalamus
Preoptic Region Habenular Nucleus
Preoptic Periventricular Nucleus Medial Habenular Nucleus
Medial Preoptic Nucleus Lateral Habenular Nucleus
Supraoptic Region Pineal Gland
Suprachiasmatic Nucleus
Anterior Nucleus MESENCEPHALON
Supraoptic Nucleus Limbic Midbrain Area
Paraventricular Nucleus Median Raphe Nucleus
Tuberal Region Dorsal Raphe Nucleus
Dorsomedial Nucleus Ventral Tegmental Area
Ventromedial Nucleus Dorsal Tegmental Nucleus
Infundibular (Arcuate) Nucleus Periaqueductal Gray
Components
Componentsof
ofthe
theLimbic
LimbicSystem
System
Limbic Lobe

Limbic Lobe Classification of Cortex


Gyrus Fornicatus
Isocortex - Neocortex
Cingulate Gyrus
Parahippocampal Gyrus Allocortex
Entorhinal Area Archicortex
Isthmus Hippocampus
Hippocampal Formation Dentate Gyrus
Hippocampus Proper Paleocortex
(Ammon’s horn)
Olfactory Bulb
Dentate Gyrus
Subiculum Piriform Lobe
Hippocampal Rudiments Mesocortex (Juxtallocortex,
Fasciolar Gyrus Periallocortex, Mesallocortex)
Indusium Griseum Cingulate Gyrus
Hippocampal Formation

Hippocampus Proper
(Ammon’s horn,
Cornu Ammonis)

Dentate Gyrus
Subiculum
Hippocampal Formation

1. uncus
2. dentate gyrus
3. fornix
4. corpus callosum
5. fasciolar gyrus
6. hippocampal sulcus
7. parahippocampal
gyrus
8. collateral sulcus
Hippocampal Formation (Lateral View)
Hippocampal Formation (Medial View)
Hippocampal Formation (Medial View)
Hippocampal Formation - Archicortex

Archicortex - 3 layers

Hippocampus Proper
Molecular Layer
Pyramidal Layer
Polymorphic Layer
Dentate Gyrus
Molecular Layer
Granular Layer
Polymorphic Layer
Subiculum
Transitional type between hippocampal archicortex and entorhinal paleocortex
Classification of Cortex

Isocortex
6 layered cortex
homotypical and
heterotypical isocortex

Allocortex
Paleocortex
olfactory cortex
- piriform lobe
Archicortex
3 layered cortex
- hippocampal formation
Archicortex - lamination by Ramon y Cajal

7.
7. stratum
stratum
moleculare
moleculare
6.
6. stratum
stratum
lacunosum
lacunosum
5.
5. stratum
stratum
radiatum
radiatum
4.
4. stratum
stratum
pyramidalis
pyramidalis
3.
3. stratum
stratum oriens
oriens
2.
2. Alveus
Alveus
1.
1. ependyma
ependyma
Hippocampus
Ammon’s horn
(Cornu Ammonis)
Subdivision
Subdivision of
of Cornu
Cornu Ammonis
Ammonis (Hippocampus
(Hippocampus proper)
proper)

-- by
by differences
differences in
in cell
cell morphology,
morphology,
relative
relative development
development andand connections
connections
-- subdivision
subdivision by
by Lorente
Lorente de
de Nó
Nó (1934)
(1934)

CA1
CA1
CA2
CA2
CA3
CA3
CA4
CA4 -- Area
Area Dentata
Dentata
Fascia
Fascia Dentata:
Dentata: outer
outer 22 layers
layers
of
of area
area dentata
dentata Rafael Lorente de Nó
Hippocampal Formation

Dentate
Dentate gyrus,
gyrus, CA1,
CA1, CA2,
CA2, CA3,
CA3, CA4,
CA4, subiculum
subiculum
Division of Hippocampal Formation

4. Fascia dentata
3. CA4
2. CA3, CA2
1. CA1
6. Subiculum
20. Presubiculum
21. Parasubiculum
22. Entorhinal cortex
Development
Developmentof
ofthe
theHippocampus
Hippocampus

1. dentate gyrus 2. hippocampus proper 3. subiculum

HS: hippocampal sulcus TH: temporal horn


Hippocampal Formation

Hippocampus Proper
CA1
CA2
CA3
CA4 - Area Dentata

Dentate Gyrus
Subiculum
Prosubiculum
Subiculum
Presubiculum
Parasubiculum
Connections of Hippocampal Formation

1.
1. Intrinsic
Intrinsic Connections
Connections
classic
classic trisynaptic
trisynaptic pathway
pathway
2.
2. Afferent
Afferent Connections
Connections
perforant
perforant and
and alveolar
alveolar path
path
3.
3. Efferent
Efferent Connections
Connections
fornix
fornix
Intrinsic Connections

Classic Trisynaptic Pathway


1. Entorhinal cortex (perforant path)
 dentate gyrus granular cell

2. Granular cell axon (mossy fiber)


 CA3 pyramidal cell

3. Pyramidal cell (Schaffer collateral)


 CA1 pyramidal cell
 subiculum  entorhinal cortex
Afferent Connections of
CA1 Pyramidal Neuron
stratum moleculare
- entorhinal cortex (perforant path)
stratum lacunosum
- Schaffer collateral from CA3
stratum radiatum
- commissural axon
- mossy fibers from dentate gyrus
stratum pyramidale
- basket cell
stratum oriens
- short axon cell
Afferent Connections

From
From Entorhinal
Entorhinal Cortex
Cortex
Alveolar
Alveolar Path
Path
from
from medial
medial part
part of
of EC
EC
to
to CA1
CA1 and
and Subiculum
Subiculum

Perforant
Perforant Path
Path
from
from lateral
lateral part
part of
of EC
EC
to
to CA1,
CA1, CA2,
CA2, CA3
CA3 and
and
Dentate
Dentate Gyrus
Gyrus
Entorhinal Cortex

1.
1. entorhinal
entorhinal cortex
cortex
2.
2. uncus
uncus
3.
3. fornix
fornix
4.
4. dentate
dentate gyrus
gyrus
5.
5. hippocampal
hippocampal sulcus
sulcus
6.
6. collateral
collateral sulcus
sulcus
Afferent Connections

EC
EC &
& Hippocampal
Hippocampal Afferents
Afferents
Surrounding
Surrounding Neocortex
Neocortex
from
from association
association areas
areas
-- temporal,
temporal, frontal
frontal lobe
lobe

Limbic
Limbic System
System
from
from hypothalamus,
hypothalamus,
amygdala,
amygdala, septal
septal area,
area,
anterior
anterior thalamic
thalamic nuclei,
nuclei,
and
and midbrain
midbrain limbic
limbic area
area
Efferent Connections

FORNIX
FORNIX
Alveus
Alveus
Fimbriae
Fimbriae
Crus
Crus
Commissure
Commissure
Corpus
Corpus
Column
Column
Efferent Connections

Fornix
- from pyramidal neurons of hippocampus & subiculum
- 1.2 million fibers in humans
Postcommissural Fornix – main bundle
to Mammillary Body
Anterior Thalamus
Lateral Septal Nuclei
Hypothalamus
Midbrain Tegmentum
Efferent Connections

Precommissural Fornix – smaller less compact


- mostly originated from the hippocampus proper
to Septal Nuclei
Lateral Preoptic Area
Anterior Hypothalamic Area
Nucleus of Diagonal Band of Broca
Nucleus Accumbens Septi
Medial Surface of Frontal Lobe
Caudate Nucleus and Putamen
Amygdala and Claustrum
Summary of Hippocampal Connections

Afferent Connections Efferent Connections


Entorhinal Cortex FORNIX
Alveolar Path from Pyramidal Neurons of
from medial part of EC hippocampus & subiculum
to CA1 and Subiculum Alveus, Fimbriae, Crus
Perforant Path Commissure, Corpus, Column
from lateral part of EC Precommissural Fornix
to CA1, CA2, CA3 and to Septal Nuclei,
Dentate Gyrus Nucleus Accumbens Septi
Dentate Gyrus Anterior Hypothalamic Area
Medial Surface of Frontal Lobe
Mossy fiber - CA3
Anterior Olfactory Nucleus
- Schaefer fiber (CA3-CA1)
Postcommissural Fornix
Others to Mammillary Body
Hypothalamus, Septal Nuclei, Anterior Thalamus
Substantia Innominata Hypothalamus
Midbrain Limbic Area Bed Nucleus of Stria Terminalis
Dopamine hypothesis of Schizophrenia
Hypersensitive
dopamine receptors
in limbic cortex

21st century
schizoid man
Septal
Septal Region
Region

Septum
Septal Region
Septal Area Septum pellucidum
Paraterminal (Subcallosal) Gyrus Septum Verum
Subcallosal Area (Parolfactory Area) - Septal Nuclei

Septal Nuclei
Dorsal Septal Nucleus Connections
Lateral septal Nucleus Stria Medullaris Thalami
Medial Septal Nucleus (Ch1) to medial habenular nucleus
Nucleus of Diagonal Band Medial Forebrain Bundle
of Broca (Ch2, Ch3) to hypothalamus and midbrain
Bed Nucleus of Stria Terminalis Fornix - to hippocampus
Bed Nucleus of Stria Terminalis & Diagonal Band
Anterior Commissure from amygdaloid complex
Basal Forebrain Area and Limbic Basal Ganglia

Basal Forebrain Area Limbic Basal Ganglia


Nucleus Accumbens Septi
(Ventral Striatum)
- structures deep to the afferents from
anterior perforating substance hippocampal formation
amygdaloid body
cingulate gyrus
Olfactory Tubercle
ventral tegmental area
Substantia Innominata dorsal raphe nucleus
Basal Nucleus of Meynert (Ch4) efferents to ventral pallidum
cf. Alzheimer’s disease Ventral Pallidum
Ventral Pallidum efferents to
- non cholinergic portions mediodorsal (MD) thalamus
substantia nigra
of the substantia innominata
subthalamic nucleus
- part of limbic basal ganglia amygdaloid nucleus
lateral habenular nucleus
Theodor Meynert
(1833-1892)

• Austrian Neurologist
• Professor of Neurology and
Psychiatry in Wien University
• Teacher of Sigmund Freud
• Use serial sections in histology
of nervous system

• Basal Nucleus of Meynert


in substantia innominata
• Dorsal tegmental decussation
of Meynert
Basal Forebrain Area Components

BASAL FOREBRAIN AREA (Heimer)

Ventral Striatum, Part of


Ventral Pallidum
Basal Nucleus of Meynert (Ch4)
Extended Amygdala

cf. Cholinergic Cell Groups (Mesulam)


(Magnocellular Basomedial Telencephalic Complex )
1. Ch1 (Medial Septal Nucleus)
2. Ch2 (Nucleus of Diagonal Band of Broca)
3. Ch3 (Nucleus of Diagonal Band of Broca)
4. Ch4 (Basal Nucleus of Meynert)
5. Ch5 (Pedunculopontine Tegmental Nucleus)
Amygdaloid Nuclear Complex

Connections
Amygdaloid Nuclear Complex Corticomedial Nuclear Group
Olfactory Pathway
Corticomedial Nuclear Group via Lateral Olfactory Stria
Anterior Amygdaloid Area Basolateral Nuclear Group
from Sensory Association Area
Cortical Amygdaloid Nucleus Central Nucleus
(Periamygdaloid Cortex) from brain stem visceral
Medial Amygdaloid Nucleus afferent nuclei
Lateral Olfactory Nucleus
Stria Terminalis
(Lateral Olfactory Gyrus) to bed nucleus of stria terminalis
Basolateral Nuclear Group hypothalamus, epithalamus
Lateral Nucleus amygdaloid body
Basal Nucleus Ventral Amygdalofugal Pathway
to septal nuclei, hypothalamus
Accessory Basal Nucleus thalamus (MD) and
Central Nucleus midbrain limbic area
Amygdala
L. almond

K. F. Burdach’s
term
Connections
Connectionsof
ofthe
theAmygdala
Amygdala
Extended Amygdala

Extended Amygdala Connections

Central Amygdaloid Nucleus Afferents from


Medial Amygdaloid Nucleus Intra-amygdaloid association
Bed Nucleus of fibers from basolateral
Stria Terminalis (BST) amygdaloid nucleus,
whic recieves wealth of
- bridging cell groups directly modality-specific and
interconnects amygdaloid multimodal sensory input
and bed nucleus of stria from the cerebral cortex.
terminalis (BST) Efferents to
which refered to as Hypothalamus and
Extended amygdala. Brain Stem
Amygdaloid
AmygdaloidNuclear
NuclearComplex
Complex Extended
ExtendedAmygdala
Amygdala

A. bed nucleus of
stria terminalis
B. centromedial amygdala
C. basolateral amygdala
D. cortical amygdala
E. connections between
extended amygdala
and amygdala:
sublenticular portion of
substantia innominata
1. stria terminalis
Amygdaloid
Amygdaloid Nuclear
Nuclear Complex
Complex

Emotion
& Amygdala
Autism
Dysfunction of
amygdala

Mercury
Rising
Limbic Thalamus

Thalamus, Limbic Connections


Anterior Nuclear Group Anterior Nuclear Group
Anteroventral Nucleus (AV) from mammillary body,
Anterodorsal Nucleus (AD) hippocampal formation,
Anteromedial Nucleus (AM) cingulate gyrus
Lateral Dorsal (LD) Nucleus to cingulate gyrus
Mediodorsal (MD) Nucleus subiculum
Pars magnocellularis (MDmc) MDmc
Pars parvocellularis (MDpc) from primary olfactory cortex,
Pars paralaminaris (MDpl) ventral pallidum
Pars densocellularis (MDdc) to orbitofrontal cortex
Midline Nuclei MDpc, MDpl, MDdc
Paratenial Nucleus (Pt) from substantia nigra
Reunience Nucleus (MV) limbic midbrain area
Submedial Nucleus (Sm) to medial frontal cortex
Rhomboid Nucleus (Rh) frontal eye field (area 8)
Connections of the Limbic Thalamus

Orbitofrontal
cingulate Cortex Frontal Medial
gyrus Eye Field Frontal
Gyrus

Primary
MB, HF Orbitory
SNr, SC, RF
Cortex
Epithalamus

Stria Terminalis
Epithalamus
from septal nuclei,
Habenular Nucleus hypothalamus,
Medial Habenular Nucleus anterior thalamic nucleus,
Lateral Habenular Nucleus substantia innominata,
Pineal Gland globus pallidus
Habenulointerpeduncular Tract
Diencephalon
(Fasciculus Retroflexus)
Epithalamus from medial habenular nucleus
Hypothalamus to interpeduncular nucleus
Thalamus (Dorsal Thalamus)
Subthalamus (Ventral Thalamus) from lateral habenular nucleus
to midbrain limbic area
Epithalamus and
surrounding structures
Hypothalamus

Hypothalamic Nuclear Group Tuberal Region


Dorsomedial Nucleus
Ventromedial Nucleus
MEDIAL ZONE
Infundibular (Arcuate) Nucleus
Preoptic Region
Mammillary Region
Preoptic Periventricular
Lateral Mammillary Nucleus
Nucleus
Medial Mammillary Nucleus
Medial Preoptic Nucleus
Intermediate Mammillary
Supraoptic Region
Nucleus
Suprachiasmatic Nucleus
Posterior Nucleus
Anterior Nucleus LATERAL ZONE
Supraoptic Nucleus Lateral Hypothalamic Nucleus
Paraventricular Nucleus Lateral Tuberal Nucleus
Hypothalamic Cell Group
Connections of the Hypothalamus

Major Afferent Connections Major Efferent Connections

1. Ascending Visceral Fibers 1. Descending Visceral Fibers


via dorsal longitudinal via dorsal longitudinal
fasciculus (DLF) and fasciculus (DLF) and
mammillary peduncle mammillary peduncle
from visceral afferent nuclei, to visceral efferent nuclei,
limbic midbrain area limbic midbrain area
2. Via Medial Forebrain Bundle 2. Via Medial Forebrain Bundle
from Septum, Amygdala and to Septum, Amygdala and
Paleocortex Paleocortex
3. Via Fornix 3. Via Mammillothalamic Tract
from Hippocampal Formation to Anterior Thalamic Nuclei
Hypothalamus

Functional Aspects of Hypothalamus

1. Endocrine Function
Hormones of the Neurohypophysis - ADH, Oxytocin
(Supraoptic and Paraventricular Nucleus)
Releasing and Inhibiting Hormones (Tuberal region)
- Hypothalamo-hypophyseal Portal System

2. Visceral Function (Autonomic Center of CNS)

3. Limbic Function
Sham Rage
- an affective attack
expression
- evoked by radio
stimulation of
medial hypothalamus

“Because the cat does not


direct its attack toward any
target, we regard this as
just a fragment of a normal
attack” (Delgado, 1981)
Silent Attack
- little emotional
expression
- evoked by electric
stimulation of
perifornical nuc.
of hypothalamus

“The cat moves swiftly


and with little sign of
emotion to bite the
rat’s neck and kill it”
(Siegel & Brutus, 1990)
Anorexia Karen Anne Carpenter
Nervosa (1950-1983)
Limbic Midbrain Area

Limbic Midbrain Area

Median Raphe (Superior Central) Nucleus (B6, B8)


Dorsal Raphe Nucleus (B7)
Ventral Tegmental Area (VTA, A10)
Dorsal Tegmental Nucleus (Gudden)
Interpeduncular Nucleus
Periaqueductal Gray
Olfactory Cortex (Rhinencephalon)

Olfactory Cortex (Rhinencephalon)

Prepiriform Cortex - Area 51


Gyrus Ambiens
Lateral Olfactory Gyrus
(Lateral Olfactory Nucleus)

Periamygdaloid Cortex
Cortical Amygdaloid Nucleus

Entorhinal Area (Cortex) - Area 28


Olfactory
OlfactorySystem
System-- Olfactory
OlfactoryPathways
Pathways

Modality: Olfaction
Receptor: Olfactory Cell of Olfactory Epithelium
Cranial Nerve: I (Olfactory Nerve)

1st Neuron: Olfactory neuron --- Olfactory nerve (I)


2nd Neuron: Olfactory Bulb --- Mitral & Tufted Cell
olfactory tract
olfactory striae
lateral & intermediate olfactory striae
Termination: Primary Olfactory Area (Rhinencephalon)
piriform lobe (prepiriform cortex, entorhinal cortex)
corticomedial amygdala
anterior perforated substance (olfactory tubercle)
Olfactory
OlfactorySystem
System-- Olfactory
OlfactoryPathways
Pathways

A. olfactory epithelium
B. olfactory bulb
C. nucleus of diagonal band
of Broca
D. periamygdaloid cortex
E. cirticomedial amygdala
F. entorhinal area
G. septal nuclei

I. olfactory nerve
1. olfactory tract
2. lateral olfactory stria
3. intermediate olfactory stria
4. medial olfactory stria
Olfactory
OlfactorySystem
System-- Connections
Connectionsof
ofPrimary
PrimaryOlfactory
OlfactoryCortex
Cortex

A. olfactory epithelium
B. olfactory bulb
C. nucleus of diagonal band
of Broca
D. periamygdaloid cortex
E. cirticomedial amygdala
F. entorhinal area
G. mediodorsal thalamic
nucleus (MD)
I. olfactory nerve
1. olfactory tract
2. lateral olfactory stria
3. intermediate olfactory stria
4. thalamocortical radiation
Archicortex and Paleocortex

hedgehog

tupaia
(tree shrew)

lemur
Connections
Connectionsof
of the
theLimbic
LimbicSystem
System
Olfactory
Olfactory
Cortex
Cortex
Septo-(Preoptico)-Hypothalamo-
Septo-(Preoptico)-Hypothalamo-Mesencephalic
MesencephalicContinuum
Continuum
Spinal
Limbid
Limbid Cord
Hippocampal
Hippocampal Septal
Septal Hypo-
Hypo- Midbrain
Midbrain &
&
Formation
Formation Region
Region thalamus
thalamus Area
Area Brain
Brain
Stem

Amygaloid
Amygaloid
body
body
Limbic System
Papez
PapezCircuit
Circuit

alveolar path Entorhinal


EntorhinalCortex
Cortex
perforant path (Area
(Area28)
28) cingulum

Hippocampal
Hippocampal Cingulate
Cingulate
Formation
Formation Gyrus
Gyrus

fornix
thalamocortical
radiation

Mammillary
Mammillary Anterior
Anterior
Body
Body Thalamic
ThalamicNuclei
Nuclei
mammillothalamic tract
Papez
PapezCircuit
Circuit

A. Hippocampal Formation
B. mammillary body
C. thalamic anterior nucleus
D. cingulate gyrus

1. fornix
2. mammillothalamic tract
3. thalamocortical radiation
4. cingulum
Limbic Function

Functional
Functional Aspects
Aspectsof
of Limbic
LimbicSystem
System
1.
1. Emotion
Emotion--Affective
AffectiveBehavior
Behavior
Amygdala
Amygdala
Limbic
LimbicandandPrefrontal
PrefrontalCotex
Cotex
--Desire,
Desire,Motivation,
Motivation,Goal-directed
Goal-directedBehavior
Behavior
Hypothalamus
Hypothalamusand andBrain
BrainStem
Stem
--ANS
ANS--maintenance
maintenanceof ofself
self(appetites)
(appetites)
maintenance
maintenanceof ofspecies
species(sexual
(sexualfunction)
function)
cf.
cf. Kl
Klüüver-Bucy
ver-BucySyndrome
Syndrome

2.
2. Memory
MemoryandandLearning
Learning
Papez
PapezCircuit
Circuit
cf.
cf. Korsakoff’s
Korsakoff’ssyndrome
syndrome
Behavioral changes of maintenance of self (appetites)
Behavioral changes of maintenance of self (aggression)

sham rage silent attack


Limbic Function

Kl
Klüüver-Bucy
ver-Bucy syndrome
syndrome
Monkey
Monkeyor
orcats
catswith
withbilateral
bilateraldamage
damageof
oftemporal
temporallobes
lobes
including
includingamygdala
amygdalaand
andhippocampal
hippocampalformation
formation

Fail
 Failto
todisplay
displaynormal
normalfear
fearof
ofanxiety
anxiety(docility)
(docility)
--attempt
attempttotopick
pickup
upsnakes
snakesand
andlighted
lightedmatches
matches

Regression
 Regressionto
tooral
oralstage
stage
--put
putalmost
almostanything
anythinginto
intomouth
mouth

Marked
 Markedincrease
increaseininboth
boththe
theamount
amountand
anddiversity
diversityof
of
sexual
sexualactivity
activity
Klüver-Bucy syndrome in cats: Mating behavioral changes
Heinrich Klüver Paul Clancy Bucy
(1897-1979) (1904-1992)
Limbic Function

Cognitive and
Emotional
Processes
Limbic Function

Evolution and Limbic System

- Triune Concept of MacLean

Protoreptilian Brain
(R Complex) ---- Instinct

Paleomammalian Brain
--- Limbic System ---- Emotion

Neomammalian Brain
--- Neocortex --- Analytical, Reasoning

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