1

Limbic System: The Center of

Emotions
Jlio Rocha do Amaral, MD & Jorge Martins de Oliveira, MD, PhD
This article by Jlio Rocha do Amaral, MD & Jorge Martins de
Oliveira, MD, PhD is brought to you by The Healing Center OnLine.

[Please see our other articles on the Neurobiology of Trauma on

our page: "FAQs, Essays & Literature related to Traumatic
Incident Reduction (TIR)"]

Introduction: The Three Units of the Human Brain

Throughout its evolution, the human brain has acquired three

components that progressively appeared and became superimposed,
just like in an archeological site: the oldest, located underneath and to
the back; the next one, resting on an intermediate position and the
most recent, situated on top and to the front. They are, respectively:

2
1 - The archipallium or primitive (reptilian) brain, comprising the
structures of the brain stem - medulla, pons, cerebellum,
mesencephalon, the oldest basal nuclei - the globus pallidus and the
olfactory bulbs. It corresponds to the reptile brain, also called "Rcomplex", by the famous neuroscientist Paul MacLean.
2 - The paleopallium or intermediate (old mammalian) brain,
comprising the structures of the limbic system. It corresponds to the
brain of the inferior mammals.
3 - The neopallium, also known as the superior or rational (new
mammalian) brain, comprises almost the whole of the hemispheres
(made up of a more recent type of cortex, called neocortex) and some
subcortical neuronal groups. It corresponds to the brain of the superior
mammals, thus including the primates and, consequently, the human
species.
These three cerebral layers appeared, one after the other, during the
development of the embryo and the fetus (ontogenesis),
recapitulating, chronologically, the evolution of animal species
(phylogenesis), from the lizards up to the homo sapiens. According to
Maclean, they are three biological computers which, although
interconnected, retained, each one, "their peculiar types of
intelligence, subjectivity, sense of time and space, memory, mobility
and other less specific functions".
Actually, we have three cerebral units in a single brain. The primitive
one is responsible for self preservation. It is there that the mechanisms
of aggression and repetitive behavior are developed. It is there that
occur the instinctive reactions of the so-called reflex arcs and the
commands which allow some involuntary actions and the control of
certain visceral functions (cardiac, pulmonary, intestinal, etc),
indispensable to the preservation of life. The development of the
olfactory bulbs and their connections made possible an accurate
analysis of olfactory stimuli and the improvement of answers oriented
by odors, such as approach, attack, flight and mating. Throughout
evolution, some of these reptilian functions were lost or minimized (in
humans, the amygdala and the entorhinal cortex are the only limbic
structures that connect with the olfactory system). It is also in the Rcomplex that started the first manifestations of the phenomena of
ritualism, by means of which the animal tries to define its hierarchic
position inside the group and to establish its own space in the
ecological niche
In 1878, the French neurologist Paul Broca called attention to the fact
that, on the medial surface of the mammalian brain, right underneath

3
the cortex, there exits an area containing several nuclei of gray matter
(neurons) which he denominated limbic lobe (from the Latin word
"limbus" that implies the idea of circle, ring, surrounding, etc) since it
forms a kind of border around the brain stem ( in another part of this
text we shall write more about these nuclei).
The entirety of these structures, that, years later would receive the
name of "limbic system", developed with the emergence of the inferior
( primitive) mammals. This system commands certain behaviors that
are necessary for the survival of all mammals. It gives rise and
modulates specific functions that allow the animal to distinguish
between the agreeable and the disagreeable. Here specific affective
functions are developed, such as the one that induces the females to
nurse and protect their toddlers, or the one which induces these
animals to develop ludic behaviors (playful moods). Emotions and
feelings, like wrath, fright, passion, love, hate, joy and sadness, are
mammalian inventions, originated in the limbic system. This system is
also responsible for some aspects of personal identity and for
important functions related to memory. And, when the superior
mammals arrived on the Earth, the third cerebral unit was finally
developed : the neopallium or rational brain, a highly complex net of
neural cells capable of producing a symbolic language, thus enabling
man to exercise skillful intellectual tasks such as reading, writing and
performing mathematical calculations. The neopallium is the great
generator of ideas or, as expressed by Paul MacLean, "it is the mother
of invention and the father of abstractive thought".

Theories on the Role of Brain Structures in the Formation of

Emotions
Comparison of the James-Lange and Cannon-Bard theories of emotion.

According to the James-Lange theory (red arrows), the man perceives

the frightening animal and reacts with physical (neurovegetative)
manifestations. As a consequence of such unpleasant physical
reaction, he develops fear. In the Cannon-Bard theory (blue arrows),
the frightening stimulus leads, first, to the feeling of fear which, then,
brings about the physical response.
At the end of the last century, William James proposed that a person,
after perceiving a stimulus that somehow affected him or her, endures
disturbing physiological changes such as palpitations, shortness of
breath, anxiety, etc.. It is precisely the acknowledgment of these
symptoms (by the brain) that creates emotion. Stating it in a different
way, we could say that physical sensations are the emotion.
In 1929, Walter Cannon refuted James's theory and advanced another
one, that was soon modified by Phillip Bardand and became known as
the Cannon-Bard theory which states that, when a person faces an
event that somehow affects him or her, the nervous impulse travels
straight to the thalamus where the message divides. One part goes to
the cortex to originate subjective experiences like fear, rage, sadness,
joy, etc. The other part goes to the hypothalamus to determine the
peripheral neurovegetative changes (symptoms). According to this
theory physiological reactions and emotional experience occur
simultaneously.

Papez believed that the experience of emotion was primarily

determined by the cingulate cortex and, secondly, by other cortical
areas. Emotional expression was thought to be governed by the
hypothalamus. The cingulate gyrus projects to the hippocampus, and
the hippocampus projects to the hypothalamus by way of the bundle of
axons called fornix. Hypothalamic impulses reach the cortex via relay
in the anterior thlamic nuclei. The essential error of the Cannon-Bard
theory was to consider the existence of an initial "center" for emotions
(the thalamus). Soon enough, though, in 1937, the neuroanatomist
James Papez would demonstrate that emotion is not a function of any
specific brain center but of a circuit that involves four basic structures,
interconnected through several nervous bundles : the hypothalamus
with its mamillary bodies, the anterior thalamic nucleus, the cingulate
gyrus and the hippocampus. This circuit (Papez circuit), acting in an
harmonic fashion, is responsible for the central functions of emotion
(affect), as well as for its peripheral expressions (symptoms).
More recently, Paul McLean, accepting the essential bases of Papez
proposal, created the denomination limbic system and added new
structures to circuit : the orbitofrontal and medialfrontal cortices
(prefrontal area), the parahippocampal gyrus and important subcortical
groupings like the amygdala, the medial thalamic nucleus, the septal
area, prosencephalic basal nuclei (the most anterior area of the brain)
and a few brainstem formations.

The Main Areas Involved with Emotions

It is important to stress that all these structures interconnect
intensively and none of them is the sole responsible for any specific
emotional state. However, some contribute more than others to this or

6
that kind of emotion. We shall review now, one by one, the best known
structures of the limbic system.

Amygdala and Hippocampus

Amygdala

Hippocampus

A little almond shaped structure, deep

inside the antero-inferior region of the
temporal lobe, connects with the
hippocampus, the septal nuclei, the
prefrontal area and the medial dorsal
nucleus of the thalamus. These
connections make it possible for the
amigdala to play its important role on the
mediation and control of major affective
activities like friendship, love and
affection, on the expression of mood and,
mainly, on fear, rage and aggression The
amygdala, being the center for
identification of danger, is fundamental
for self preservation. When triggered, it
gives rise to fear and anxiety which lead
the animal into a stage of alertness,
getting ready to flight or fight.
Experimental destruction of both
amygdalas (there are two of them, one in
each hemisphere) tames the animal, which
becomes sexually non-discriminative,
deprived of affection and indifferent to
danger. The electrical stimulus of these
structures elicits crises of violent
aggressivity. Humans with marked lesions
of the amygdala, loose the affective
meaning of the perception of an outside

Is particularly involved with memory

phenomena, specially with the formation
of long-term memory (the one that,
sometimes, lasts forever). When both
hippocampi (right and left) are destroyed,
nothing can be retained in the memory.
The subject quickly forgets any recently
received message. The intact hippocampus
allows the animal to compare the
conditions of a present threat with similar
past experiences, thus enabling it to choose
the best option, in order to guarantee its
own survival.

information, like the sight of a well

known person. The subject knows,
exactly, who the person is, but is not
capable to decide whether he likes or
dislikes him (or her).

Fornix and Parahippocampal gyrus

Both are important connecting pathways of the limbic system.

Thalamus and Hypothalamus

Thalamus
Lesion or stimulation of the medial dorsal and anterior nuclei of the
thalamus are associated with changes in emotional reactivity.
However, the importance of these nuclei on the regulation of emotional

8
behavior, is not due to the thalamus itself, but to the connections of
these nuclei with other limbic system structures. The medial dorsal
nucleus makes connections with cortical zones of the pre-frontal area
and with the hypothalamus. The anterior nuclei connect with the
mamillary bodies, and through them, via fornix, with the hippocampus
and the cingulate gyrus, thus taking part in the Papez's circuit.

Hypothalamus
This structure has ample connections with the other prosencephalic
areas and the mesencephalus. Lesions of the hypothalamic nuclei
interfere with several vegetative functions and some of the so-called
motivated behaviors, like thermal regulation, sexuality, combativeness,
hunger and thirst. The hypothalamus is also believed to play a role in
emotion. Specifically, its lateral parts seem to be involved with
pleasure and rage, while the median part is like to be involved with
aversion, displeasure and a tendency to uncontrollable and loud
laughing. However, in general terms, the hypothalamus has more to do
with the expression (symptomatic manifestations) of emotions than
with the genesis of the affective states. When the physical symptoms
of emotion appear, the threat they pose returns, via hypothalamus, to
the limbic centers and, thence, to the pre-frontal nuclei, increasing
anxiety. This negative feed-back mechanism can be so strong as to
generate a situation of panic. As it will be seen later on, the knowledge
of this phenomenon is very important, for clinical and therapeutic
reasons.

Cingulate gyrus

It is located in the medial side of the brain between the cingulate

sulcus and the corpus callosum (principal fiber bundle connecting the
two cerebral hemispheres). There is still much to be learned about this
gyrus, but it is already known that its frontal part coordinates smells
and sights with pleasant memories of previous emotions. This region
also participates in the emotional reaction to pain and in the regulation
of aggressive behaviour. Wild animals, submitted to the ablation of the
cingulate gyrus (cingulectomy), become totally tamed. The cutting of a
single bundle of this gyrus (cingulotomy) reduces pre-existent

9
depression and anxiety levels, by interrupting neural communication
across the Papez's circuit.

Brainstem

The brainstem is the region responsible for the "emotional reactions",

(indeed, they are just reflex answers) of inferior vertebrates, like
reptiles and amphibians. The involved structures are the reticular
formation, and the locus coeruleus, a concentrated mass of norepinephrine secreting neurons. It is important to stress that, even in
humans, these primitive structures remain active, not only as alerting
mechanisms, vital for survival, but in the maintenance of the sleepawake cycle.

Ventral Tegmental Area

In the ventral tegmental area, located in the mesencephalic part of the

brain stem, there is a compact group of dopamine-secreting neurons
whose axons end in the nucleus accumbens (mesolimbic dopaminergic
pathway). The spontaneous firing or the electrical stimulation of
neurons belonging to that region produce pleasurable sensations,

10
some of them similar to orgasm. Many people who, for a genetic error,
have a reduction of D2 (dopamine) receptors in the accumbens
nucleus, become, sooner or later, incapable to obtain gratification from
the common pleasures of life. Thus, they seek atypical and noxious
"pleasurable" alternatives, like alcoholism, cocaine addiction, impulsive
gambling and compulsion for sweet foods. Certain brainstem
structures, like the nuclei of the cranial nerves, stimulated by impulses
coming from the cortex and the striatum (a subcortical formation), are
responsible for the physiognomic : expressions of anger, joy, sadness,
tenderness, etc.

Septum

The septal region lies anteriorly to the thalamus. Inside it, one finds
the centers of orgasm (four for women and one for men). This
area has been associated with different kinds of pleasant sensations,
mainly those related to sexual experiences.

Prefrontal area

11
This area comprises the entire non-motor anterior region of the frontal
lobe. It underwent a great deal of development during the evolution of
mammals. It is specially large in man and in some species of dolphins.
It does not belong to the traditional limbic circuit, but its intense bidirectional connections with thalamus, amygdala and other subcortical
structures, account for the important role it plays in the genesis and,
specially, in the expression of affective states. When the pre-frontal
cortex suffers a lesion, the subject looses his sense of social
responsibility as well as the capacity for concentration and abstraction.
In some cases, although consciousness and some cognitive functions,
like speech, remain intact, the subject can no longer solve problems,
even the most elementary ones. When pre-frontal lobotomy
was used for treatment of certain psychiatric disturbances,
the patients entered into a stage of "affective buffer", no
longer showing any sign of joy, sadness, hope or despair. In
their words or attitudes, no traces of affection could be
detected.

Affective States
Humans display the largest web of connections between
the prefrontal area and the traditional limbic structures.
Perhapas that is why they present, among all species, the
greatest variety of feelings and emotions Although some
signs of affection can be perceived in birds, the limbic system only
began to evolve, in fact, after the first mammals, being practically nonexistent in reptiles, amphibians and all other preceding species.
Paul MacLean uses to say that "it is very difficult to imagine a lonelier
and more emotionally empty being than a crocodile". Two behaviors,
with affective connotation, that appeared with mammals (birds also
display them, but less intensely) deserve to be emphasized because of
their peculiarity :
1 - The intense and long-lasting care and nursing of females towards
their offsprings.
2 - A playful mood. The more evolved the mammal, the more
accentuated are these behaviors.
Ablation of important parts of the limbic system of any animal causes it
to loose, totally, both motherly affection and ludic interest.

12
And the evolution of mammals bring us to mankind. Certainly, our
hominid ancestor could already establish differences between the
sensations he experienced in distinct occasions, such as being at his
cave polishing a stone or a bone, running after a weaker animal,
running away from a stronger one, hunting a female of his species etc.
With the development of language, particular names were given to
these sensations, allowing their definition and communication to other
members of the group. Since there exists an important subjective
component, difficult to be communicated, even today there is no
uniformity concerning the best terminology to be used, in order to
designate, specifically, many of these sensations.
Therefore, the words affect, emotion and feeling are used
interchangeably and imprecisely, almost like synonyms. However, we
think that each of these words deserves a precise definition, for the
sake of their etymology and because of the physical and mental
reactions they cause.
Affect (from the Latin affectus meaning to afflict, to shake, to touch)
could be defined as "a grouping of physic phenomena manifesting
under the form of emotions, feelings or passions, always followed by
impressions of pleasure or pain, satisfaction or discontentment , liking
or disliking, joy or sorrow".
Curiously, there is a worldwide trend to consider as affect (as well as
its derivatives, like affection, affectuous, etc.) only the positive
impressions. Thus, when I say : "I feel affection for that girl", I'm
expressing love or tenderness, never anger or fear. Counterwise,
emotions and feelings can be used to name both positive and negative
phenomena: "she has good feelings; I've had painful emotions".
According to Nobre de Melo, affect denominates, generically, events
experienced as emotions or feelings. Emotions (from Latin emovere
meaning moving, displacing) are, as its etymology suggests, manifest
reactions to those affective conditions that, due to their intensity,
move us to some kind of action. Confronting the opinion of several
authors, we can say that emotions are characterized by a sudden
disruption of the affective balance.
Almost always, they are short episodes, with slight or intense,
concomitant or subsequent, repercussions upon several organs, that
can set up partial or total blocking of logical reasoning. This can
provoke, in the affected subject, a high degree of psychic and
behavioral loss of control. Conversely, feelings are seen as affective
states with a longer duration, causing less intensive experiences, with

13
much fewer repercussions upon organic functions and lesser
interference on reasoning and behavior.
Exemplifying: love, fear and hate are feelings; passion, fright and
anger (or wrath) are emotions. There also exist two other well
characterized conditions that are, in a certain way, inserted in the
context of affective life (since, depending on the intensity of the affect,
these two conditions may result from the affective state or, sometimes,
can be mistakenly considered as an emotional manifestation).
We are talking about mood disturbances (represented by depressions
or maniacal euphoria) and the lessening of mental relaxation
associated to the alert reaction (represented by anxiety). During
centuries, philosophers, physicians and psychologists studied the
phenomena of affectivity, questioning their origin, their role upon
psychic life, their action favoring or hindering adaptation, their
neurophysiological concomitants and their neuroendocrine substratum.
Affective manifestations would have as their ultimate cause, the
capacity of living matter to answer to stimuli acting upon it. There are
two classical and antagonistic theories upon the subject. The first one,
supported by Darwin and his followers, state that affective reactions
are innate patterns designed to orient behaviour in order to promote
the adaptation of a being to its environment, thus guaranteeing its
survival and that of its species.
The organic disturbances, that may go together with the process,
would only be a consequence of physiological nature. On the contrary,
others, like William James, state that, facing a given real or imaginary
stimulus, the organism would react with a series of muscular and
visceral neurovegetative changes.
The perception of such changes would give origin to the corresponding
affective states. More recently, Lehmann proposed a third theory that
suggests a compromise between the two classical ones. According to
him, affect is a complex phenomenon, initiated by a central process, as
result of internal or external causes. It manifests itself as a change in
the "I", releasing reflex facial movements and producing manifold
organic alterations.
The more the bodily symptoms increase, the more mobilizing becomes
the affect, until it evolves into an emotion. This statement finds clinical
support in the treatment of patients with performance phobias. When
facing situations they fear (speaking before audiences, for instance)
these patients present palpitations, sweating, difficult breathing, etc..
Beta blocking agents, that do not cross blood brain barrier (therefore
devoid of influence upon brain centers) act only peripherally, blocking

14
the neurovegetative symptoms and, by "emptying" anxiety, facilitate
the control of the phobic process.
Divergent are also the opinions about the relationship between
affective states and reason. Some philosophical and religious schools
consider the affective aspects of personality as inferior, negative or
sinful, in need of control and domination by reason. Claparde, in a
paper bearing the title "Feelings and emotions", defines emotions as
useless, unadaptative and harmful phenomena, true remains of
ancestral reactions. Just the opposite of feeling, that would be useful,
allowing human beings to estimate the value of things to which they
must adapt and differentiating the useful from the noxious.
Quoting the author : "Observation shows how unadaptive are
emotional phenomena. Emotions happen precisely when adaptation is
hindered by any motive . "The analysis of corporal reactions in
emotions shows that the subject does not enact adaptive movements
but, on the contrary, reactions that resemble indefinite primitive
instincts... "Far from being the psychic aspect of an instinct, emotion
represents a confusion of this instinct ". Contrariwise, other authors
consider affective reactions as factors favoring adaptation and survival,
inducing some behaviours and inhibiting others. In their opinion, even
intense emotions, evaluated as disruptive by others, could favour
survival.
This is so because their disruptiveness is a selective one : while some
actions are abolished, others, more favorable, are allowed to occur. We
believe that, within certain limits, the affective participation reinforces
the cognitive component, giving more flavour to the day-to-day
experiences and facilitating adaptive behaviors. Nevertheless, when
above these limits, emotions hamper reasoning and, when below them,
as stressed by Damasio in "Descarte's error", affectivity becomes
scarce, thus impoverishing the quality of life.

About the Authors:

Jlio Rocha do Amaral, MD Teacher of clinical pharmacology,
anatomy and physiology. Medical
Manager of Merck S/A Indstrias
Qumicas (pharmaceutical and
chemical industries). Redactor of
didactic manuals on anatomy,

Jorge Martins de Oliveira, MD, PhD. Full Professor and Master of UFRJ (Rio de
Janeiro). Associate Professor of UFF. Scientific
Coordinator. Coordinador and Director of the
Department of Neurosciences of the Institute of
Human Being (RJ). Fellow in Research by Saint
Vincent Charity Hospital, Cleveland, USA. Full

15
physiology and pharmacology used
by Merck S/A. Editing supervisor of
the following scientific publications:
Senecta, Galenus and Sinapse.
Redactor of clinical trials and
protocols since 1978. Assistant
coordinator of courses on Oxydology
sponsored by the Human Being
Institute and UNIGRANRIO
(University of Great Rio). Head of
Psychiatric Service. Neurosciences
Department. The Human Being
Institute. Co-author of the book
"Principles of Neurosciences".
Email: j.r.amaral@mls.com.br

Member of Brazilian Academy of Military

Medicine. Member of the Brazilian Academy of
Writers Physicians. Graduated by Superior School
of War (ESG).
Email:jmartins@rio.nutecnet.com.br