International Review of Psychiatry

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Neurobiology of emotions: an update

Vanderson Esperidião-Antonio, Marilia Majeski-Colombo, Diana Toledo-

Monteverde, Glaciele Moraes-Martins, Juliana José Fernandes, Marjorie
Bauchiglioni de Assis, Stefânia Montenegro & Rodrigo Siqueira-Batista

To cite this article: Vanderson Esperidião-Antonio, Marilia Majeski-Colombo, Diana Toledo-

Monteverde, Glaciele Moraes-Martins, Juliana José Fernandes, Marjorie Bauchiglioni de Assis,
Stefânia Montenegro & Rodrigo Siqueira-Batista (2017): Neurobiology of emotions: an update,
International Review of Psychiatry, DOI: 10.1080/09540261.2017.1285983

To link to this article: http://dx.doi.org/10.1080/09540261.2017.1285983

Published online: 25 May 2017.

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INTERNATIONAL REVIEW OF PSYCHIATRY, 2017
http://dx.doi.org/10.1080/09540261.2017.1285983

ARTICLE

Neurobiology of emotions: an update

Vanderson Esperidi~ao-Antonioa, Marilia Majeski-Colombob, Diana Toledo-Monteverdeb,
Glaciele Moraes-Martinsb, Juliana Jose Fernandesb, Marjorie Bauchiglioni de Assisb, Stef^ania Montenegroc
and Rodrigo Siqueira-Batistac,d,e
a
Departamento de Medicina e Enfermagem, Universidade Federal de Viçosa (UFV), Viçosa, Brazil; bCurso de Graduaç~ao em Medicina,
 ~aos (UNIFESO), Teresopolis, Brazil; cN
Centro Universitario Serra dos Org ucleo de Estudos em Ci^encias Medicas, Faculdade Din^amica do
Vale do Piranga (FADIP), Ponte Nova, Brazil; dDepartamento de Medicina e Enfermagem, Universidade Federal de Viçosa (UFV),
Viçosa, Brazil; ePostgraduate Program of Bioethics, Ethics and Public Health (PPGBIOS), Universidade Federal do Rio de Janeiro (UFRJ),
Rio de Janeiro, Brazil

ABSTRACT KEYWORDS
The ‘nature’ of emotions is one of the archaic themes of Western thought, thematized in differ- Limbic system; emotions;
ent cultural manifestations – such as art, science, philosophy, myths and religion –, since Ancient neurosciences; neurobiology
times. In the last decades, the advances in neurosciences have permitted the construction of
hypotheses that explain emotions, especially through the studies involving the limbic system. To
present an updated discussion about the neurobiology of processes relating to emotions –
focusing (1) on the main neural structures that relate to emotions, (2) the paths and circuits of
greater relevance, (3) the implicated neurotransmitters, (4) the connections that possess neurove-
getative control and (5) the discussion about the main emotions – is the objective of this pre-
sent article.

Introduction of a thinking substance (res cogitans) that is com-

The study of emotions, in terms of knowledge, is
rather archaic in Western culture (Espiridi~ao Antonio
et al., 2008). As a matter of fact, turning back on
ancient Greece, it is possible to find in the concept of
pathos (pahos) the idea of passion, understood in the
hellenic philosophical ambit as the root of evil and
the unhappiness of man (Reale, 1999), and should
therefore be moderated and, perhaps, dominated. It is
noted in this horizon, for example, the platonic con-
ception of the tripartite soul, being that the portions
that correspond to emotions are the mortal ones –
lustful and irascible – and the rational soul is the
immortal (divine) portion (Plat~ao, 1977; Siqueira-
Batista & Schramm, 2004).
From this prospective, the complex emotion/reason
relation has become a recurrent motto in the thoughts
of the different philosophers, which formulated con-
ceptions, in the most various manner, to explain the
origins and the role of different emotions in the
human condition. This was precisely the case of
french thinker Rene Descartes, that, in his
Metaphysical Meditations, arrives at the conceptions
pletely separated from the ‘worldly substance’ (res
extensa), empowering, in this manner, a genuine
schism between body and mind (Descartes, 2016). To
Descartes, the res cogitans belongs to reason and to
thought, while emotions pertain to the body (res
extensa), characterized as confused and non credible
in relation to the true contents (Descartes, 2016). On
the other hand, the philosopher Baruch of Spinoza
conceived that the ‘thinking substance and the exten-
sive substance are the same substance, sometimes
understood as one attribute and other times as
another’ (Spinoza, 1997: Part II, Prop. VII) (Spinoza,
2010). In these terms, reason and emotion – and the
organic constitution itself – would belong to the same
nature (Spinoza, 2010).
From a genuine philosophical problem, the rela-
tions between the body and mind and between reason
and emotion began being investigated in the theoret-
ical ambit of other sciences – such as psychology, psy-
choanalysis, biology and medicine –, from the second
half of the XIX century and the beginning of the XX
century. What marks this period is the scientific inter-

This is an updated English version of the paper: Esperidiao-Antonio V. et al. (2008). Neurobiologia das emoç~oes. Rev. Psiquiatr. Clín., 35, 55–65.
ß 2017 Institute of Psychiatry and Johns Hopkins University
2 ~
V. ESPERIDIAO-ANTONIO ET AL.
est appointed to the cognitive processes, which
includes the mental activities related to the acquisition
of knowledge and connected to reasoning and mem-
ory. This elucidation is explained by the greater com-
mensurability of cognition, leading to the
development of the so called ‘cognitive revolution’.
From then on, numerous investigations that culmi-
nated in the proposition of the mechanisms involved
in perception, in attention and in memory were real-
ized. The few authors that focused on emotions, con-
ceived them in a ‘segmented’ manner, treating the
‘emotional circuits’ as separate events that are inde-
pendent from other neural activities.
More recently – since the development of new spe-
cialized research techniques in neurophysiologiy and
neuroimaging (Billot et al., 2017; Brooks et al., 2016;
Esperidi~ao Antonio & Majeski-Colombo, 2012;
George, 2016) –, there has been a broadening of inter-
est in the study of neural bases of the processes
involved in emotions, as of the characterization and
the investigation of the limbic system (LS). It is
known, based on different results, that there is a deep
integration between the emotional, cognitive and
homeostatic processes, in a manner in which its iden-
tification would be of great value for the better under-
standing of the physiological response of the organism
when the individual is faced with various situations.
Therefore, connections with diverse nervous circuits,
which, through their neurotransmitters promote
physiological responses that relate the organism to the
environment (somatic nervous system) as well as to
the innervation of visceral structures (visceral nervous
system), that are important for the maintenance of
the internal environment’s stability (homeostasis)
(Yang et al., 2007).
Despite these advances, the possibility of treating
issues regarding emotions scientifically has been
greatly discussed. With the development of the neuro-
sciences, it is postulated that, like the perception and
the action, emotion is related to distinct cerebral cir-
cuits. Furthermore, emotions are generally accompa-
nied by autonomic, endocrine and skeletal motor
responses – that depend on the subcortical areas of
the nervous system –, which prepare the body for
action (Felger & Miller, 2014; Kandel, 2000; Lanotte
et al., 2005). In fact, it is believed that science will be
capable of explaining biological aspects related to
emotions, but not what emotion is: this remains a
predominantly philosophical issue. Based on these
premisses, the objective of the present article is to dis-
cuss the current aspects of the neurobiology of emo-
tions – with the LS as a starting point –, therefore
punctuating the important relation between the emo-
tional processes and the autonomic nervous system,
emphasizing its interference in the neurovegetative
control.
Historical background: from the limbic system
to the system of emotions
Since the Renaissance, period marked, among other
aspect, by a substantive progress of human anatomical
studies – morphological investigations, typified by a
fruitful composition between science and art (Barreto
& Oliveira, 2004), have been developed, which permit
a description of different organic structures, including
those of the brain and the other constituents of the
central nervous system (CNS) and peripheral nervous
system (PNS). This context was one of the factors
responsible for the subsequent interest in compre-
hending the cerebral processes related to cognition
and emotions, theme that has won special impulse
since the XVIII and XIX century (Esperidi~ao Antonio
& Majeski-Colombo, 2012; Gazzaniga, Ivry, &
Mangun, 2013), especially after the works of Gall,
Broca and Papez.
Franz Joseph Gall described the morphology of the
brain and the main nervous structures, which facili-
tated a significant advance in differentiating the
important cerebral portions, characterizing some of its
specific functions, becoming known as ‘the author of
the true anatomy of the brain’. The first ‘mapping’ of
cerebral functions was proposed by Pierre Paul Broca,
fulfilled through the observation of patients with cere-
bral damaged. Broca identified the limbic lobe
(limb ¼ margin), which comprises a ring composed by
continued cortical structures situated in the medial
and inferior face of the brain (Damasio, Grabowski,
Frank, Galaburda, & Damasio, 1994).
A large impulse for the establishment of hypotheses
about the neural bases of emotions took place from
the description of the Phineas Gage case, a worker
who survived an incident that provoked severe cere-
bral damage consequent of a serious work related
injury (Damasio et al., 1994; Harlow, 1868). After an
accidental explosion, Gage had his head pierced by a
metal bar, which penetrated through the left maxillary
region, deeply reaching his frontal lobe ipsilaterally.
Gage did not lose conscience, despite his frontal lobe
being injured. After a few weeks, the worker wished
to return to work but did not receive permission from
his employer due to great behavioral changes. Before
the incident, Gage was a capable and efficient worker
with a well balanced mind, seen as an astute, intelli-
gent and talented man. After the morbid event, he

became indecisive demonstrating indifference, falsity,
disloyalty and sloppiness. He also showed impatience
and was unable to establis any plan for the future
(Lent, 2010).
The interest of the comprehension of the mental
and cerebral processes can also be identified in the
pioneer investigations developed, in the last century –
by the viennese physiologist and the psychologist
Sigmund Exer, by the psychoanalyst Sigmund Freud
and by the french doctor Israel Waynbaum –, in
which knowledge about the neural networks and pos-
sible structures that would compose the emotional cir-
cuits was being illustrated (Harlow, 1868). In this
movement, temporarily situated in the transition of
the XIX century to the XX century, the first neuro-
psychological theories of emotions were proposed,
highlighting the conceptions of William James and
Carl Lange – to whom the subjective emotional
experience would be consequent of the physiological
and behavioral manifestations (in other words, if one
is happy it is because one smiles … ) – as well as the
ideas of Walter Cannon and Phillip Bard – that for-
mulated the the CNS caused both the subjective
experience and the physiological and behavioral mani-
festations (Cannon, 1927).
Substantive advance for the comprehension of the
neurobiological phenomenons related to emotion was
reached by Joseph Papez, American anatomists that
shifted the gaze of an emotional centers perspective,
substituting it for a conception of systems. In fact, ini-
tially it was believed that the limbic lobe described by
Broca would be related to smell, but Papez demon-
strated that its different portions were united and
coordinated within itself, forming a circuit, which
included the cingulate cortex, the hippocampus and
the anterior nuclei of the thalamus (Cannon, 1927).
Further experimental evidences permitted the revi-
sion of structures pertaining to the circuit proposed
by Papez, therefore emerging the concept of the lim-
bic system (LS) – resuming the term created by Broca
in describing the limbic lobe –, which would involve
the structures related to emotions (Cosenza, 1990;
Papez, 1937). The LS began being characterized as a
neural circuit related to the emotional responses and
to the motivational impulses, having already been
included in his design, structures such as the hypo-
thalamus, amygdala, nuclei of the bases, prefrontal
area, cerebellum and septum (the hippocampus, ini-
tially inserted, does not appear to have decisive par-
ticipation in the neural mechanisms of emotions,
having role in the consolidation of memories, includ-
ing those with emotional content, therefore being
INTERNATIONAL REVIEW OF PSYCHIATRY 3
related – although not pertinent – to the LS)
(Cannon, 1927; Dror, 2001). In a similar manner, the
group of anterior nuclei of the thalamus is not con-
firmed as a substantive element in the neurobiology
of emotions. In point of fact, there still does not exist
a perfect accord on the components of the LS.
Although the denomination ‘limbic system’ is still
used to designate components involved in the cerebral
circuits of emotions, this categorization has been
receiving critiques in many degrees. For example, the
inclusion of diverse anatomical structures in its
design, without clear agreement between authors; in
fact, the majority of the investigators include, in the
LS, the cingulate gyrus, the parahippocampal gyrus,
the amygdala, the hypothalamus and the area of the
septum. Other structures such as the cerebellum, thal-
amus, prefrontal area and hippocampus are not
always said to be pertinent to the LS, although these
elements possess direct relations to the emotional
processes and the autonomic responses (Guyton &
Hall, 2011; Machado, 2013; Papez, 1937; Pergher,

Grassi-Oliveira, De Avila, & Stein, 2006). This fact
indicates that there is no criteria broadly accepted to
decide upon the constitution of the LS (Fontenelle, de
Oliveira-Souza, & Moll, 2015). Furthermore, the idea
itself of a single system of emotions has been placed in
check ever since the identification of different circuits
and areas of the CNS that correlate to the disparate
states called emotions. Thus, in the present manu-
script, the structures consensually recognized by the
different authors, as well as those that directly relate
to emotions with be included in this discussion about
the LS.
Therefore, substituting – or amplifying – the idea
of the LS, a conception of the systems of emotions
(Espiridi~ao Antonio et al., 2008; Esperidi~ao Antonio &
Majeski-Colombo, 2012), that house the disparate cir-
cuits and neural networks that correlate to the typified
states such as emotion, should be proposed.
Structures related to emotions
The systems of emotions (Espiridi~ao Antonio et al.,
2008; Esperidi~ao Antonio & Majeski-Colombo, 2012)
– at least as they have been understood recently –
appear to be organized in a network; more pro-
nounced morphofunctionally regulatory components
do not exist in these systems, in other words, all of
the elements play regulatory roles similar to each
other (LeDoux, 2003). Therefore, it is understood that
these systems depend on the integration of their com-
ponents in a complex, non-hierarchical manner,
which still needs to be further explained (Papez, 1937;
4 ~
V. ESPERIDIAO-ANTONIO ET AL.
Figure 1. Structures of the limbic system. Neuroanatomical
image prepared by the authors.
Table 1. Structures that compose the circuit of Papez.
Structures Comments
Cingulate gyrus It is related to depression, anxiety, and aggression; in
humans, mental retardation is observed in cases of
injury to this structure. It assists in determining the
contents of memory, with a significant increase in
their activity when people lie (Guyton & Hall, 2011).
Parahippocampal It is related to memory storage; Injurious processes
gyrus located there produce isolated retrograde amnesia,
with preservation of the capacity to store new
explicit memories (Gazzaniga et al., 2013).
Hypothalamus This structure, for Papez, would constitute the central
segment of the limbic system, relating to the vari-
ous encephalic areas. Both hypothalamic stimula-
tion and inhibition often have profound effects on
the behavior and emotions of animals, including
Homo sapiens sapiens. Stimulation of the lateral
hypothalamus induces thirst, hunger and increases
the general level of activity of the animal, some-
times leading to fury and/or fighting. The stimula-
tion of the ventromedial nucleus causes a contrary
situation, that is, a sensation of satiety, reduction of
food intake and tranquility (Pergher et al., 2006).
The stimulation of periventricular nuclei usually
causes fear and punishment reactions. The sex
drive can be stimulated mainly in the anterior and
posterior portions of the hypothalamus.
Thalamus The functions are mainly related to sensitivity, motor,
emotional behavior and activation of the cerebral
cortex (Papez, 1937).
Hippocampus The hippocampus has important functions related to
behavior and memory (Guyton & Hall, 2011).
People undergoing bilateral hippocampal removal
can access learned memory but can not learn any
new information. This area is also integrated with
decision making, because when the hippocampus
interprets a neuronal signal as important, it is prob-
ably stored in memory (Pergher et al., 2006).
Recently the relationship between the hippocampus
and the immune system has been demonstrated,
identifying that its integrity is fundamental for the
normality of the immune response, as well as the
interaction of memory with the levels of interleukin
1 alpha (IL-1a) and IL-2. The hippocampus is not
currently considered a crucial part of the neurobio-
logical systems of emotions.
Table 2. Structures related to the emotions and that are not
included, originally, to the circuit of Papez.
Structures Comments
Amygdala It is activated in situations with marked emotional signifi-
cance, such as aggressive or sexual meeting; Is also
related to emotional learning and the storage of affect-
ive memories. In addition, the amygdala is responsible
for the formation of the association between stimuli
and rewards.
Septum The septum is related to anger, pleasure and neurovege-
tative control. It has been shown in animals that bilat-
eral involvement of the septal area causes “septal
anger”, characterized by emotional hyperactivity, fer-
ocity and anger in situations that do not usually
change animal behavior. Changes in blood pressure
and respiratory rate may be observed when the septal
area is stimulated (Yang et al., 2007). Self-stimulation
experiments performed on rats allowed the localization
of “pleasure areas” in the brain; Among the most fre-
quently stimulated areas are the septal area and the
regions covered by the medial forebrain bundle. This
hypothesis was, in part, confirmed in experiments with
human patients(Gazzaniga et al., 2013).
Prefrontal The prefrontal area has been considered the core of the
cortex personality (Gazzaniga et al., 2013). There is still much
speculation around this region, but through the inter-
pretation of experimental and clinical data, it is noted
that this structure participates in decision making and
in the adoption of behavioral strategies more appropri-
ate to the physical and social situation; In addition, it
seems to be related to the capacity to follow ordered
sequences of thoughts and the modalities of control of
the emotional behavior (Benoit, 2008; Guyton & Hall,
2011).
Cerebellum Nowadays, it has been recognized that this organ has
functions broader than purely motor, acting in several
cognitive processes. Cerebellar damage is associated
with dysfunctions in executive tasks, learning, proced-
ural and declarative memory, language processing and
visual and spatial functions, as well as dysfunctions in
personality, affection and cognition. The hypothesis
deriving from the anatomical model is that the rupture
of the neural circuit that connects the cerebellum with
the associative and paralimbic areas prevents cerebellar
modulation of related cognitive functions, causing
alterations in the subsystems and producing conduct
deficits (LeDoux, 2003). It was proposed a scheme of
the different types of non-motor activity, which could
be modulated by different cerebellar regions. In the
case of cognition and emotion, the earliest cerebellar
regions, such as the nodule-lobe, the worm, the fasti-
gial nucleus and the globose nucleus are described,
which can be considered equivalent to a limbic cere-
bellum, being responsible for the mechanisms
Primitives of preservation, as manifestations of struggle,
emotion, sexuality and, possibly, emotional memory
(LeDoux, 2003). The cerebellar lateral hemispheres and
the mandibular and emboliform nuclei seem to be
responsible for the modulation of thought, planning,
strategy formulation, learning, memory and language,
characteristics that are only identified in mammals. In
this way, the cerebellum has been considered a power-
ful coordinator, capable of contributing to both motor
and sensory and cognitive abilities, thanks to the con-
nections it establishes with brain regions responsible
for performing these functions (Guyton & Hall, 2011).
Berridge, 2004). Figure 1 and Table 1 present the
components of the LS, as proposed by Papez. Table 2
exposes additional structures that integrate with the
system of emotions (Espiridi~ao Antonio et al., 2008;
Esperidi~ao Antonio & Majeski-Colombo, 2012).

Although there is no precise definition of the
neural circuit involved in the complex system of emo-
tions (Espiridi~ao Antonio et al., 2008; Esperidi~ao
Antonio & Majeski-Colombo, 2012), some neuronal
pathways can be described, in a didactic fashion, with-
out loosing sight of the fact that they are ultimately
functionally integrated. These paths will be discussed
in the latter context of the different emotions.
The neural bases of emotions
Pleasure and reward
The most ‘primitive’ emotions that are also well
studied by neurophysiologists – with the purpose of
establishing their relations with cerebral functioning –
are the sensations of rewards (pleasure, satisfactions)
and punishment (disgust, aversion), each one having
been characterized as their own encephalic circuit.
The ‘center of reward’ is primarily related to the
medial prosencephalic bundle – in the lateral and
ventromedial nuclei of the hypothalamus –, having
connections with the septum, amygdala, some areas of
the thalamus and the ganglions of the base (Lent,
2010; Dror, 2001; Pergher et al., 2006). On the other
hand, the ‘punishment center’ is described as being
located in the central grey area that surrounds Sylvius’
cerebral aqueduct in the midbrain, extending to the
periventricular zones of the hypothalamus and thal-
amus, being related to the amygdala and the hippo-
campus (Dror, 2001) as well as the medial portions of
the hypothalamus and the lateral portions of the teg-
mental areas of the midbrain (Ballone, 2002).
For some researchers, the sensations of please can
be distinguished by the facial expressions and attitude
of the animal after exposure to a hedonic stimulus;
these expressions are maintained even in anencephalic
individuals, suggesting that the ‘center of please’ must
extend to the cerebral trunk (LeDoux, 2003). It is
believed that afferent emissions of the accumbens
nucleus towards the lateral and ventral hypothalamus,
the globus pallidus and the structures connected in
this same cerebral region are involved in the hedonic
cerebral circuits (LeDoux, 2003).
In investigations with experiment animals (rats), it
was demonstrated that stimuli in the septum area,
controlled by the animal, entailed a stimuli of recur-
rent deflagration of the stimuli, indicating a possible
correlation with the triggering of pleasure (Esperidi~ao
Antonio & Majeski-Colombo, 2012; Gazzaniga et al.,
2013). This conclusion, however, is not so simple: for
example, in the 1960s, Robert Heath applied 17 elec-
trodes on the encephalon of a patient in order do
INTERNATIONAL REVIEW OF PSYCHIATRY 5
discover the location of severe epilepsy. What the
researcher was able to observe was that the stimula-
tion in specific areas, according to the implantation
of the electrodes, generated different types of sensa-
tions. The patient in this case stimulated more fre-
quently a site in the medial thalamus that, although
it provoked an irritable sensation, caused an immi-
nent sensation of evoking a memory, making him
repeat the procedure in order to bring back the
memory to mind, in other words, with the objective
of obtaining a rewards with repeated stimulation
(Ballone, 2002).
Later studies on simians demonstrated participation
of the medial proencephalic bundle on the appetitive
stimuli, being able to characterize a certain expect-
ation of pleasure. This bundle and its integrated
regions (ventral tegmental area, hypothalamus, accum-
bens nucleus, anterior cingulate cortex and prefrontal
cortex) compose the circuit called mesolimbic system.
Dopamine appears to be fundamental in mediating
the rewards’ effects. Dopaminergic neurons project
from the ventral tegmental area of the midbrain to
many areas of the encephalon through the medial
prosencephalic bundle. Furthermore, drugs that cause
chemical dependency increase the effectiveness of
dopamine and provoke its release in the accumbens
nucleus, demonstrating the role of this neurotransmit-
ter in the mechanisms of reward and/or pleasure
(Ballone, 2002). However, the investigations carried
out until now continue obtaining ambiguous results,
especially in the ambit of pharmacological studies –
which indicate that the dopamine’s agonists increase
the auto-stimulation rate, while the antagonists
decrease this rate – and the lesion studies, that show
that the section of the medial prosencephalic bundle,
por example, does not provoke significant reduction
of the auto-stimulation, as expected (Ballone, 2002).
Happiness
The experiences nuanced by happiness – response to
the identification of facial expressions of joy, and with
the visualization of pleasant images and/or the induc-
tion of memories of happiness, sexual pleasure and
successful competitive stimulation – provokes the acti-
vation of basal ganglions, including the ventral corpus
striatum and the putamen (Bear, Connors, &
Paradiso, 2008; Niculescu, Schork, & Salomon, 2010).
The recent revision published by Machado Cantilino
(Bogert et al., 2016) demonstrated that the postitive
emotions – including joy and happiness – relate with
the reduction of activity in the right prefrontal cortex
and the increase of activity in the left prefrontal
6 ~
V. ESPERIDIAO-ANTONIO ET AL.
regions, as well as the association with the increment
of activities in the regions of the cingulate gyrus,
inferior and middle temporal gyrus, amygdala and
ventral striatum (Bogert et al., 2016). Furthermore, it
is important to remember that the basal ganglions
receive vast innervation from the dopaminergic neu-
rons of the mesolimbic system – intimately related to
the generation of pleasure – and the dopaminergic
system of the ventral striatum (Machado & Cantilino,
in press; Suardi, Sotgiu, Costa, Cauda, & Rusconi,
2016). Dopamine acts independently – utilizing opioid
and GABAergic receptors on the ventral striatum, the
amygdala and on the orbitofrontal cortex – fact
related to the affective sates (such as sensorial pleas-
ure), while other neuropeptides are involved in the in
the generation of the sensation of satisfaction through
the homeostatic mechanisms (Li et al., 2016).
Neuroanatomical descriptions of the lesions of the
cerebral-pontocerebellar pathways in individuals with
pathological laughter and crying suggest that the cere-
bellum is a structure involved in the association
between the execution of laughter and crying and the
cognitive and situacional context in question; in fact,
when such structure is injured there in an incomplete
transition of information, provoking inadequate
behavior in its context (Burgdorf & Panksepp, 2006).
Empathy and compassion
The investigations directed towards empathy and
compassion have produced interesting results, in neu-
roscientific terms. The two concepts are similar, but
do not completely overlap. In fact, empathy refers to
the ‘union or fusion with all beings or objects (consid-
ered animated)’ (Parvizi, Anderson, Martin, Damasio,
& Damasio, 2001); in a distinct manner, it is also con-
ceived as ‘an action of unconditional welcoming’, with-
out judgment (Abbagnano, 2012).
Based on these considerations, empathy and com-
passion can be characterized as phenomenons intim-
ately related to emotional control and the capacity of
relating with others. They are generally considered as
pertaining to the Homo sapiens sapiens specie, even
though studies with other animals species have dem-
onstrated the existence of neural circuits that can be
correlated to empathetic behaviors (Siqueira-Batista &
Schramm, 2009).
Subsequently, research that used functional neuroi-
maging techniques pointed to the existence of intri-
cate neural circuits – involving the premotor cortex,
the parietal lobe and the supplementary motor area –
which are activated by preforming a task or visualiz-
ing the same task performed by someone else, which
corroborates to the hypothesis of the existence of an
ideomotor component in empathy (Meyza, Bartal,
Monfils, Panksepp, & Knapska, 2016). In this sense,
Enzy et al. (Enzi, Amirie, & Br€ une, 2016) (Sampaio,
Camino, & Roazzi, 2009) highlight that empathy is
associated to the activity in a complex neural network,
including the anterior insula, the anterior and middle
cingulate cortex and the lateral prefrontal cortex. The
role of the prefrontal cortex appears to be fundamen-
tal, consequentially deserving attention from research-
ers – especially the ventrolateral prefrontal cortex and
the dorsolateral prefrontal cortex (Sampaio et al.,
2009; Enzi et al., 2016) –, highlighting that the trans-
cranial stimulation of continued current of the latter
can ‘modulate’ empathy (Ferrari, 2014).
Another important research frontier refers to the
neuroendocrine aspects, highlighting that there
appears to be a correlation between empathy and the
action of certain hormones, such as oxytocin and
vasopressin (Wang, Wang, Hu, & Li, 2014).
Fear
The sensations of fear and anger are related to the
existing significant relations between the amygdala
and the hypothalamus (Uzefovsky et al., 2015). The
amygdala is responsible for the detections, generation
and maintenance of the emotions related to fear, as
well as those related to the recognition of fearful
facial expressions and the coordination of the appro-
priate responses to threat and danger (Bear et al.,
2008; Mobbs, Hagan, Dalgleish, Silston, & Prevost,
2015; De Gelder, Snyder, Greve, Gerard, &
Hadjikhani, 2004). An injury of the amygdala in
humans produces a reduction in the emotionality
and the capacity of recognizing fear. On the other
hand, the stimulation of the amygdala can lead to a
state of vigilance or of increased attention, anxiety
and fear (Ballone, 2002; H€ oistad & Barbas, 2008; Lee,
Milton, & Everitt, 2006).
Since the initial descriptions of the LS, realized by
Papez, it is believed that the hypothalamus plays a
crucial role between the subcortical structures
involved in the processing of emotions. Currently, it
is recognized that the projections of the amygdala to
the cortex contribute to the recognition of experienc-
ing fear and other cognitive aspects of the emotional
process (Fontenelle et al., 2015).
The amygdala is a structure that acts preforms as
an essential link between the areas of the cerebral cor-
tex, receiving information from all of the sensorial
systems. These, in turn, project in a specific form to
the amygdaloid nuclei permitting the integration of

information coming from many cerebral areas,
through excitatory and inhibitory connections from
the cortical and subcortical pathways (Graham,
Devinsky, & LaBar, 2006). The basolateral nuclei are
the main entrance ways of the amygdala, receiving
sensorial and auditory information; however the ven-
tral amygdalofugal pathways and the terminal stri-
atum establish a connection with the hypothalamus,
permitting the triggering of fear (Ballone, 2002). The
terminal striatum is related to the release of stress
hormones form the pituitary and adrenal gland during
conditioning (Fontenelle et al., 2015).
Sensory inputs to the amygdala are received by the
lateral nucleus. The auditive inputs come form the
auditive thalamus and the auditive cortex and arrive
to the lateral nucleus of the amygdala, stimulating it
in the processes of conditioned fear (as described
below). This fact is confirmed by functional magnetic
resonance studies (fMRI) on humans, in which, dur-
ing conditioning, activity in the amygdala and corre-
lating activity in the thalamus were observed
(Fontenelle et al., 2015; Williams et al., 2006). In add-
ition to the auditive cortex and thalamus, ventral
areas of the hypothalamus project to the basolateral
and basomedial nuclei of the amygdala, and in case of
injury to these areas, there is interference in the gen-
eration of the conditioning. The central nucleus of the
amygdala is responsible for the interface of the motor
system, consequentially, injuries to this nucleus reveal
alterations in the expression of responses to condi-
tioned fear (Fontenelle et al., 2015).
The role of the amygdala in the triggering of fear
was better studied in the 1970s and thee 1980s, when
the technique of pavlovian fear conditioning was
described. This technique consisted of offering and
emotionally neutral stimulus, such as the emission of
an audible pitch (conditioned stimulus), and associat-
ing it to an aversive stimulus, such an electric shock
(unconditioned stimulus). After the repeated applica-
tion of theses associated stimuli, it was noted that the
conditioned stimulus was able to provoke responses
typically observed in the presence of danger, such as
defense behavior (flight or fight response), activation
of the autonomic nervous system (alterations in blood
flow and cardiac frequency), neuroendocrine
responses (release of pituitary and adrenal hormones),
among others (Fontenelle et al., 2015). Situations such
as the exposure to loud and subtle sounds, great
heights and large visual stimuli that are not identified
– that emerge suddenly from the superior part of the
field of vision –, produce the so called unconditioned
fear, present in many animals. In the human species,
INTERNATIONAL REVIEW OF PSYCHIATRY 7
the unconditioned fear can be produced, for example,
by darkness. The conditioned fear, or learned fear, is
caused by the majority of stimuli, that become
‘warnings’ that a situation is threatening and can hap-
pen again (Gazzaniga et al., 2013). After this discov-
ery, it was possible to associate these behaviors to the
amygdala, once the injuries in this structure interfered
in the acquisition and the expression of the conditions
fear (Fontenelle et al., 2015).
The investigation of the amygdala through imaging
exams, such as positron emission tomography (PET)
and fMRI, allowed concluding the this structure is
activated even when the analyzed individual is not
directly submitted to a situation that provokes fear. In
addition, it was observed that the amygdala is not
activated only processes that involve the sensation of
fear, but also during more positive situations, such as
during the recognition of happy facial expressions,
leading to the conclusions that the amygdala is
involved in the response to importante emotional
stimuli, regardless of its pleasant or unpleasant con-
text (Bear et al., 2008). It should also be highlighted
that the amygdala is related to the ability to ‘read’
ocular movements expressed in facial mimics, during
significant emotional manifestations (Gonzalez-Villa
et al., 2016; Dejean et al., 2015).
In order to learn the conditions fear, the pathways
that transmit information from the stimulus converge
at the lateral nucleus of the amygdala, from the the
information goes to the central nucleus. This, in turn,
establishes a connections with the hypothalamus and
the periaquedutal gray matter of the cephalic trunk,
evoking, at last, somatic motor responses (LeDoux,
2003; Ballone, 2002). The participation, in this pro-
cess, of the basolateral nucli of the amygdala could be
confirmed from the observation, in investigational
studies of the NMDA receptors of the brain – recep-
tors associated to the acquisition, reconsolidation and
extinction of memories. It was verified that the amyg-
daloid nuclei were the primary location for the actions
of drugs that act upon the NMDA receptors, in other
words, potentiating (agonist drugs) or extinguishing
(antagonist drugs) the memory associated to the con-
ditioned fear (H€ oistad & Barbas, 2008).
In addition to the important role of the amygdala
in generating fear, this process can also appear to be
dependent on the serotonergic, noradrenergic and
central GABAnergic system, to the extent that it is
known that the mechanism of action of the anti-
depressant and anxiolytic pharmaceuticals are depend-
ent on the interferences of the habitual conduction of
these pathways, when altering the concentration in its
8 ~
V. ESPERIDIAO-ANTONIO ET AL.
receptors (Jacobs, Renken, Aleman, & Cornelissen,
2012).
The role of the temporal lobe in the neurobiology
of fear should also be highlighted. injuries in this
location produce alteration in social and emotional
behavior of animals, such as the acquisition of a
docile posture by wild and ferocious animals, lost of
fear, extreme curiosity, quick forgetfulness, the ten-
dency to place everything in mouth and extremely
intense sexual impulse (Kluver-Busy syndrome).
Although similar disturbances are rare in human
being, the affected people react in a similar manner
to the simians – condition characterizes by apathy,
lethargy and emotional insensibility (Pergher et al.,
2006).
Anger
Anger is basically manifested by aggressive behaviors,
almost always intense, which depend on the involve-
ment of diverse structures and organic systems in
order to be expressed (Lara & Akiskal, 2006).
Furthermore, this behavior also allows for variations
according to the stimulus thats evokes it.
One of the first structures associated to anger was
the hypothalamus, due to studies realized in the
1920s, in which was the description of manifestations
of anger in unsuitable situations, after total removal
of the telencephalon. However, this same behavior
was not observed when the injury extended to the
posterior half of the hypothalamus, leading to the
conclusion that the posterior hypothalamus would be
involved in the expression of anger and aggressive-
ness, while the telencephalon would mediate inhibi-
tory effects on this behavior (Ballone, 2002).
Generally, two behaviors, classically studied in ani-
mals, can be describes: a predatory aggression, that has
an objective the obtaining of food, and affective
aggression, whose purpose is exhibition to other sur-
rounding animals or females. During the 1960s, John
Flynn identifies that these aggressive behaviors were
provoked by the stimulation of specific areas of the
hypothalamus, located respectively on the medial and
lateral hypothalamus (Beyer, M€ unte, G€ ottlich, &
Kr€amer, 2014).
The typical behavior of predatory aggression can be
verified after stimulating the lateral hypothalamus,
which possesses many outputs in the ventral tegmen-
tal area through medial prosencephalic bundle. On
the other hand, the section of this neuronal bundle
does not totally eliminate such behavior, indicating a
possibility that the hypothalamus is not the only
structure associated to the generation of this
behavioral pattern. Affective aggression, in turn, is
provoked by stimulus from the periaquedutal gray
matter by the lateral hypothalamus through the dorsal
longitudinal fascicle (Ballone, 2002).
Fear, such as anger, is and emotion related to the
functions of the amygdala, in result of connections to
the hypothalamus and other structures (Flynn, 1967).
The electric stimulation of the basolateral nucli of the
amygdala activate the hypothalamus and the nuclei of
the encephalic trunk through the ventral amygdalofu-
gal pathway, producing the typical behavior of affect-
ive aggression. On the other hand, the stimulation of
the corticomedial nuclei provoked inhibitory outputs
to the hypothalamus through the temrinal striatum,
reducing predatory aggression (Ballone, 2002).
A study was carried out with the objective of identi-
fying the cerebral areas involved in the processing of
anger, utilizing Fmri (Zhao, Sun, Chen, & Yang, 2016).
The stimulus received by the participants were furious
or neutral voices, simultaneously, and they would
choose which one to listen to. The results of the studies
showed that the right amygdala and the bilateral super-
ior temporal grooves responded to the recognition of
anger, regardless of when the voice that denoted anger
was chosen or not; therefore, the orbitofrontal cortex
and the cuneus (in the medial occipital cortex) showed
greatest activation when the furious voice was chosen
rather than when it was discarded, indicating possible
association of these areas with neural processing of the
recognition of fear (Zhao et al., 2016). In addition to
this study, there are description of the association of the
injuries of the orbitofrontal cortex – especially in the
context of the influence of emotion in the process of
decision making (Lara & Akiskal, 2006) – to inappro-
priate behaviors such as impulsiveness, anger, little
expression of happiness and characteristics of double
personality disorder (Sander et al., 2005). Furthermore,
in another investigation utilizing imaging exams
(fMRI), bilateral activation of the amygdala was dem-
onstrated during visualization of scared faces, while the
observation of the figures that denoted infuriated facial
expression provokes stimulation of only the right
amygdala. Such fact, besides agreeing with the study
described previously (Zhao et al., 2016), makes it pos-
sible to reinforce that the amygdala is associated to the
perception of threatening facial expressions and that its
connection to the other cortical and subcortical struc-
tures permit a response to this threat (Berlin, Rolls, &
Iversen, 2005; Fitzgerald, Angstadt, Jelsone, Nathan, &
Phan, 2006; Suslow et al., 2006).
In addition to the structural components, there are
studies that involve the participation of

neurotransmitter in the modulation of anger and
aggression. Serotonin is one of the neurotransmitters
involved in this regulation, which can easily suggest,
once the location of the serotonergic neurons on the
encephalic trunk’s raphe, the medial prosencephalic
bundle, the hypothalamus and in other associated lim-
bic structures (Jacobs et al., 2012). This association
can be reinforced by studies realized with knockout
mice for the 5-HT1B receptors. These specific recep-
tors are localized on the nuclei of the rafe, the amyg-
dala, the periaquedutal grey matter and the ganglions
of the base (Ballone, 2002).
Anger appears to be mainly modulated by the
accumbens nucleus and through the dopaminergic and
glutamatergic systems, once dopaminergic antidepres-
sants and psychostimulants are potentialize fear and
the antipsychotics and mood stabilizers can have
depressant effects on anger (Jacobs et al., 2012).
Fight or flight reactions
The hypothalamic projections to the regions of the
brainstem – highlighting the nucleus tractus solitarius –
are possibly responsible for the direct connections
between the hypothalamus and the autonomic nervous
system (ANS). In addition to the output pathways, the
vagus nerve (CN X), one of the main elements of the
ANS (parasympathetic portion), represent an important
afferent component, activating superior cerebral areas:
its input projections ascend to the forebrain through
the parabraquial nucleus and the ceruleus locus, con-
necting directly to all of the levels of the forebrain
(hypothalamus, amygdala and thalamic regions that
control the insula and the orbitofrontal and prefrontal
cortex) (Majeski-Colombo, Viana, Pinto, Esperidi~ao-
Ant^ onio, & Siqueira-Batista, 2006; Porges, 2003). In
this way, the vagus nerve is highlighted in its participa-
tion in the integrated response (cognition-emotion),
because, besides stimulating the encephalic areas, it
produces direct physiological organic reactions (the
vagal inputs will determine specific responses, inhibi-
tory or excitatory, in various tissues and bodily systems)
and also indirect, through the parallel stimulation of
other nerves that originate from the next vagal center of
the central nervous system.
The ANS is directly involved in the so called
‘situations of fight and/or flight’ and immobilization
(Atzori et al., 2016). These events are intrinsically
related to a mechanism of neuroception, that is char-
acterized by the capacity of an individual of acting
according to his/her perception of his/her environ-
ment as safe or dangerous. This perception can be
given by the tone of voice or the movements and
INTERNATIONAL REVIEW OF PSYCHIATRY 9
facial expressions of a person or animal that the indi-
vidual communicates with (Majeski-Colombo et al.,
2006; Porges, 2003).
Every time a person perceives an environment as
‘safe’, he/she has inhibitory mechanisms that act on
the limbic structures that control the fight or flight
behaviors, including the lateral and dorsomedial
regions of the periquedutal grey matter (Kahle, Miller,
Lopez, & Hastings, 2016). This mechanism can be
exemplified by the neural projections of the fusiform
gyrus and superior temporal sulcus to the amygdala
(more precisely the central amygdaloid nucleus), that
inhibit it. In this manner, the amygdala does not play
its normal role, in other words, the stimulation of
these pathways in the periaquedutal grey matter.
Concomitantly, after the processing of all of the infor-
mation, the motor cortex (where the frontal areas are
highlighted) command the activation of the cortico-
bulbar pathways of the medulla (source nuclei of the
V, VII, IX, X and XI cranial nerves), that activate the
somatomotor components (muscles of the face and
head) and visceromotor (heart, bronchial tree) of the
physiological mechanisms for social contact (Majeski-
Colombo et al., 2006).
On the other hand, every time a person perceives
the environment as ‘dangerous’, the amygdala will be
free to trigger excitatory stimuli upon the lateral and
dorsolateral region of the periaquetudal grey matter,
that then stimulate the pathways of the pyramidal
tract, producing fight or flight responses. In addition,
there are cases where the person responds to such
situation as if she/he were paralyzed; this response
happens due to the stimulation of the region that is
ventrolateral to the cerebral aqueduct (Sylvius), that
also stimulates the neural pathways of the lateral cor-
ticospinal tracts (pyramidal). It is interesting to
emphasize that these reactions occur parallel to the
sympathetic autonomic response, by nerves that ori-
ginate from the paravertebral ganglions, and sympa-
thetic (Nerves III, VII, IX and X), as well as its sacral
portion, represented by the sacral nerves from S2 to
S4, thus allowing the innervation of blood vessels and
smooth muscles of organs from different systems in
all of the organism. In fight or flight situation there is
an elevation of the cardiac frequency and of the sys-
temic arterial blood pressure; however, in situations of
immobilization, intense bradycardia and systemic
arterial hypotension occurs.
Sadness
Sadness and depression can be seen as ‘poles’ of a
same process – the first is considered ‘physiological’,
10 ~
V. ESPERIDIAO-ANTONIO ET AL.
and the second, ‘pathological’ – therefore being
related to in neurophysiological terms (Bondarenko,
Guimar~aes, Braga, & Nalivaiko, 2016; Gelenberg,
2010). The description of the correlation between
emotional dysfunctions and damage to the neurocog-
nitive functions are more and more frequent. As a
matter of fact, depression associates to deficits in stra-
tegic areas of the brain, including the limbic regions.
Different investigations demonstrated that the real-
ization of activities that evoke this sentiment are
related to the activation of the central areas, such as
the inferior and medial occipital gyri, the fusiform
gyrus, the lingual gyrus, the postero-medial and
superior gyri and the dorsal amygdala, also highlight-
ing the participation of the dorsomedial prefrontal
cortex (Barrett & Janata, 2016), which was reinforced
in recent studies utilizing musical evocation
(Niculescu et al., 2010). Furthermore, in normal indi-
viduals it was observed, through positron emission
tomography (PET) that the induction of sadness
related to: 1) the activation of the limbic regions –
subgenual portion of the cingulate gyrus and the
anterior insula –; 2) cortical deactivation – right pre-
frontal cortex and inferior parietal cortex –; and 3)
reduction of glucose metabolism in the prefrontal cor-
tex (Goldin et al., 2005). In the same manner, import-
ant activation of the subcallosum cingulate cortex
(especially in the anterior subgenual/ventral cingulate
region) was identified after induction of sadness in
the studied individuals; however, in patients with clin-
ical depression, hypometabolism or hypofusion was
noted in the subcallosum cingulate cortex (Bear et al.,
2008).
With the objective of evaluating the neurochemical
systems involved in the emotional processes, Zubieta
et al. (Machado-Vieira, Bressan, Frey, & Soares, 2005)
realized a study on humans, observing that, through
the stimulation and maintenance of a saddened state
and deactivation of the neurotransmitters of the anter-
ior rostral cingulate gyrus, the ventral pallidum, the
amygdala and the inferior temporal cortex was devel-
oped (Machado-Vieira et al., 2005). There is a correl-
ation between the increase of the rate of negative
feelings and a reduction of the rate of positive feel-
ings, confirming the role of mu-opioid receptors in
the regulation of the affective human experiences.
Emotion and reason
The relation between emotion and reason represent
archaic themes in Western Culture, motivating
important philosophical reflections in the last 25 cen-
turies (Zubieta et al., 2003). In scientific terms, an
important addition to the investigation in this field
was experienced from the 1990 up until today, pro-
ducing promising results (Siqueira-Batista, Batista, &
Schramm, 2012). In fact, it is generally recognized
that the information that reaches the brain, that com
from different sources, goes through a certain trajec-
tory where it is processed. Then it is directed to the
the limbic and paralimbic structures, through the
neural pathways related to emotions – for example,
the Papez circuit –, or through other connections, to
acquire emotional significance, directing itself, con-
tinuously, to certain regions of the cerebral cortex,
permitting that decisions be made and actions trig-
gered – processes related to autonomy (Figure 2) – a
function that is generally dependent of the frontal or
prefrontal cortex (Benoit, 2008; Damasio & Carvalho,
2013; Posner & Raichle, 1994; Siqueira-Batista &
Schramm, 2008).
For the most part, images certainly provoke activa-
tion of the visual occipital cortex (occipital and fusi-
form gyri), however, the amygdala can also receive a
substantial amount of stimuli coming from the tem-
poral areas associated to vision, participating in the
formation of memories through the hippocampal or
the striatum circuits (Fontenelle et al., 2015). Such
Figure 2. Is it possible to propose a “neuroanatomy” of deci-
sion-making involving the prefrontal cortex – perhaps a neuro-
biology of autonomy? Recent investigations have corroborated
to this hypothesis. In this context, the use of reason would be
initiated medially by the action of the anterior cingulate cortex
(executive attention), which has as a function focusing the per-
ceptual and cognitive attention, modulating the activity of the
corresponding areas. The dorsolateral areas of the prefrontal
cortex would be responsible for the comparison of new and
old information. The ultimate adjustment – taking into account
the objective of teach individuals and the social contexts –
would be realized by an area that is not illustrated, the ventro-
medial prefrontal cortex. Neuroanatomical image prepared by
Rodrigo Siqueira-Batista (UFV and FADIP) and Vanderson
Esperidi~ao Antonio (UFV), inspired by Posner and Raichle
(1994).

fact occurs due to the special role of the amygdala in
the processing of relevant visual emotional informa-
tion, signaling fear and aversion or other evidences.
The activation of the amygdala can be mainly
involved in the emission of an alert for upcoming
threats obtained from the perception by the cortex
(Bear et al., 2008).
The integration of the affective load to the cogni-
tive processes probably occurs in the orbitofrontal
cortex complex (OFC)/ventromedial prefrontal cortex
(VMPFC) (Abu-Akel, 2003; Siqueira-Batista &
Schramm, 2012; Siqueira-Batista & Schramm, 2013).
In fact, the sensorial impressions (such as vision,
hearing, and other somatosensory information) con-
verge through the PFC to the VMPFC, from where
the synthesized information is taken to the dorsome-
dial and infero-lateral prefrontal cortex regions for
decision making. Injuries in the VMPFC cause dam-
age to the decision-making capacity, generally charac-
terized by the inability to adopt strategies of
appropriate behavior to the consequences of actions
taken, leading to impulsiveness. The VMPFC and the
OFC maintain important relation to the amygdala and
both contribute to decision-making, although the
mechanisms by which it occurs are distinct. It is
believed that these cortical regions receive input from
the amygdala – which represent the motivational
value of the stimuli – integrating themselves and pro-
moting an evaluation of future behavior that will be
adopted (Gamond & Cattaneo, 2016; De Martino,
Kumaran, Seymour, & Dolan, 2006).
Although the amygdala does not establish direct
connection to the prefrontal lateral cortex, it com-
municates with the anterior cingulate and orbital
cortex, which are involved in memory circuits, mak-
ing the justification of some authors possible that
the amygdala participates in the modulation of mem-
ory and in the integration of emotional and cogni-
tive information, possibly attributing emotional load
to them, enabling the transformation of subjective
experiences to emotional experiences (Fontenelle
et al., 2015; Gamond & Cattaneo, 2016). Injuries of
the medial rostral prefrontal cortex lead to the
inappropriate expression of emotions during social
behaviors and damage to the personally beneficial
decision-making process (Bear et al., 2008; Siqueira-
Batista & Schramm, 2013).
Studies show that the executive processes are medi-
ated by the frontal lobe, particularly the anterior cin-
gulate cortex (ACC) and the prefrontal cortex (PFC).
According to these investigation, the medial PFC is
involved in associating the cognitive aspect to the
INTERNATIONAL REVIEW OF PSYCHIATRY 11
emotional one, being responsible for the evaluation
and/or cognitive interruption of emotions (Bear et al.,
2008; Gamond & Cattaneo, 2016; Morawetz, Bode,
Baudewig, & Heekeren, 2016).
More recently, it was described that the ACC and
the dorsolateral prefrontal cortex (DLPFC) hold the
neural bases of the performance of concurrent tasks
of central execution. The results show that the cogni-
tive processes are modulated by the ACC of the
same hemisphere. The ACC is probably positioned
in a way that connects information within the brain;
this way, it includes reciprocal connections that are
important to the motor system and the emotional
system. The DLPFC participates in the executive sit-
uations, such as coordination of concurrent cognitive
processes and selective attention to information of
relevant tasks. It is postulated that general human
intelligence is a result of a functional integration
between the alert system based on the ACC and the
executive system based on the DLPFC (Kondo,
Osaka, & Osaka, 2004).
Another important structure in integrating emo-
tion/reason is the insula, which is activated during
the induction of memories of experienced moments,
which provoke a specific sensation, whether it be
happiness, sadness, pleasure, anger, or any other,
for specific stimuli (Billot et al., 2017). However, the
insula is not activated when the same sensation is
provoked in the same individual through a film, sug-
gesting that the structure is involved in aspects of
evaluation, experimentation or expression of an emo-
tion generated internally (Bear et al., 2008).
Based on what was discussed, it is possible to
the consider that decision-making becomes directly
dependent of the emotional association realized by
the individual when he/she experiences certain daily
situations. Furthermore, the use of this acquired
information depends on motor responses – such as
running, smiling or eating – and autonomic
responses, such as the elevation or reduction of the
cardiac frequency or increased intestinal peristalsis.
Such autonomic responses are directly influenced
by the hypothalamus and it, in turn, acts
through processing all of the information that
reaches the brain.
Final considerations
The correlation of (1) different emotions and (2)
interactions between emotional states and cognitive
processes – including decision-making – to neural
structures (with special emphasis on its
12 ~
V. ESPERIDIAO-ANTONIO ET AL.
anatomofunctional characteristics) is still the reason
for interrogation and motivation for scientific investi-
gation (Gainotti, 2012; Hacker, 2009; Kondo et al.,
2004), as this present article sought to delineate.
Different stimuli (inputs) – thermic, tactile, visual,
auditive, olfactory and of visceral nature (such as
alterations in blood pressure) –, reach different parts
of the CNS through neuronal pathways involving per-
ipheral receptors and nerves. Adequate responses
(outputs) to these same stimuli are programmed in
certain cortical areas, which range from simple cir-
cuits – involving few segments – to complex neural
pathways, demanding functional refinement from
each and every part.
The circuits related to emotions are located in
various regions of the encephalon, possessing
numerous connections to the cortex, subcortical area
(matter), its nuclei and infratentorial structures –
belonging to the brainstem and cerebellum. The
relation to the brainstem is further highlighted,
facilitating synapses to the reticular matter, nuclei
such as the red nucleus, the ambiguous nucleus and
the nuclei that make up the cranial nerves, high-
lighting those that make up the III pair (oculo-
motor nerve), VII pair (facial nerve), IX pair
(glossopharyngeal nerve) and the X pair (vagus
nerve), belonging to the cranial portion of the para-
sympathetic nervous system. From then on, a stimu-
lus is directed to the cerebellum and the spinal
cord, distributed by the spinal nerves to bodily seg-
ments and to the sympathetic nervous system by
the segments from T12 to L1 (thoracolombar) and
to the parasympathetic nervous system from S2 to
S4 (sacral part). This is a panoramic view of the
biological integration between emotions and the
neurovegetative control.
In spite of these already constructed knowledges –
which allow for the proposal of different, but inte-
grated, systems of emotions (Espiridi~ao Antonio et al.,
2008), highly amplifying a reductionist/localizationist
view of the superior mental functions –, there is still
a long way to go to acquire a better understanding
of the fundamental neurobiological mechanisms
related to – perhaps determinants of – emotions,
path that can supposedly be capable of bringing
man/woman to the understanding of her/his own
condition.
Disclosure statement
The authors report no conflicts of interest. The
authors alone are responsible for the content and writing of
this article.
Funding
The authors are grateful to CNPq and FADIP for financial
support.
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