The 

limbic system, also known as the paleomammalian cortex, is a set of brain structures located

on both sides of the thalamus, immediately beneath the medial temporal lobe of
the cerebrum primarily in the forebrain.[1]
Its various components support a variety of functions including emotion, behavior, long-term
memory, and olfaction.[2]
The limbic system is involved in lower order emotional processing of input from sensory systems and
consists of the amygdaloid nuclear complex (amygdala), mammillary bodies, stria medullaris, central
gray and dorsal and ventral nuclei of Gudden.[3] This processed information is often relayed to a
collection of structures from the telencephalon, diencephalon, and mesencephalon, including
the prefrontal cortex, cingulate gyrus, limbic thalamus, hippocampus including the parahippocampal
gyrus and subiculum, nucleus accumbens (limbic striatum), anterior hypothalamus, ventral
tegmental area, midbrain raphe nuclei, habenular commissure, entorhinal cortex, and olfactory
bulbs.[3][4][5]

Structure[edit]

Anatomical components of the limbic system

The limbic system was originally defined by Paul D. MacLean as a series of cortical structures
surrounding the boundary between the cerebral hemispheres and the brainstem. The name "limbic"
comes from the Latin word for the border, limbus, and these structures were known together as
the limbic lobe.[6] Further studies began to associate these areas with emotional and motivational
processes and linked them to subcortical components that were then grouped into the limbic system.
[7]

Currently, it is not considered an isolated entity responsible for the neurological regulation of
emotion, but rather one of the many parts of the brain that regulate visceral autonomic processes.
[8]
 Therefore, the set of anatomical structures considered part of the limbic system is controversial.
The following structures are, or have been considered, part of the limbic system:[9][10]

 Cortical areas:
o Limbic lobe
o Orbitofrontal cortex: a region in the frontal lobe involved in the process of decision-making
o Piriform cortex: part of the olfactory system
o Entorhinal cortex: related to memory and associative components
o Fornix: a white matter structure connecting the hippocampus with other brain structures,
particularly the mammillary bodies and septal nuclei
 Subcortical areas:
 o Septal nuclei: a set of structures that lie in front of the lamina terminalis, considered a
pleasure zone
o Hippocampus and associated structures: play a central role in the consolidation of new
memories
o Amygdala: located deep within the temporal lobes and related with a number of emotional
processes
o Nucleus accumbens: involved in reward, pleasure, and addiction
 Diencephalic structures:
o Hypothalamus: a center for the limbic system, connected with the frontal lobes, septal
nuclei, and the brain stem reticular formation via the medial forebrain bundle, with the
hippocampus via the fornix, and with the thalamus via the mammillothalamic fasciculus;
regulates many autonomic processes
o Mammillary bodies: part of the hypothalamus that receives signals from the hippocampus
via the fornix and projects them to the thalamus
o Anterior nuclei of thalamus: receive input from the mammillary bodies and involved in
memory processing

Function[edit]
The structures and interacting areas of the limbic system are involved in motivation, emotion,
learning, and memory. The limbic system is where the subcortical structures meet the cerebral
cortex.[1] The limbic system operates by influencing the endocrine system and the autonomic nervous
system. It is highly interconnected with the nucleus accumbens, which plays a role in sexual
arousal and the "high" derived from certain recreational drugs. These responses are heavily
modulated by dopaminergic projections from the limbic system. In 1954, Olds and Milner found
that rats with metal electrodes implanted into their nucleus accumbens, as well as their septal nuclei,
repeatedly pressed a lever activating this region.[11]
The limbic system also interacts with the basal ganglia. The basal ganglia are a set of subcortical
structures that direct intentional movements. The basal ganglia are located near the thalamus and
hypothalamus. They receive input from the cerebral cortex, which sends outputs to the motor
centers in the brain stem. A part of the basal ganglia called the striatum controls posture and
movement. Recent studies indicate that if there is an inadequate supply of dopamine in the striatum,
this can lead to the symptoms of Parkinson's disease.[1]
The limbic system is also tightly connected to the prefrontal cortex. Some scientists contend that this
connection is related to the pleasure obtained from solving problems.[citation needed] To cure severe
emotional disorders, this connection was sometimes surgically severed, a procedure
of psychosurgery, called a prefrontal lobotomy (this is actually a misnomer). Patients having
undergone this procedure often became passive and lacked all motivation.[12]
The limbic system is often incorrectly classified as a cerebral structure,[citation needed] but simply interacts
heavily with the cerebral cortex. These interactions are closely linked to olfaction, emotions, drives,
autonomic regulation, memory, and pathologically to encephalopathy, epilepsy, psychotic symptoms,
cognitive defects.[13] The functional relevance of the limbic system has proven to serve many different
functions such as affects/emotions, memory, sensory processing, time perception, attention,
consciousness, instincts, autonomic/vegetative control, and actions/motor behavior. Some of the
disorders associated with the limbic system and its interacting components are epilepsy and
schizophrenia.[14]

Hippocampus[edit]
Location and basic anatomy of the hippocampus, as a coronal section

The hippocampus is involved with various processes relating to cognition and is one of the best
understood and heavily involved limbic interacting structure.
Spatial memory[edit]
The first and most widely researched area concerns memory, particularly spatial memory. Spatial
memory was found to have many sub-regions in the hippocampus, such as the dentate gyrus (DG)
in the dorsal hippocampus, the left hippocampus, and the parahippocampal region. The dorsal
hippocampus was found to be an important component for the generation of new neurons, called
adult-born granules (GC), in adolescence and adulthood.[15] These new neurons contribute to pattern
separation in spatial memory, increasing the firing in cell networks, and overall causing stronger
memory formations. This is thought to integrate spatial and episodic memories with the limbic
system via a feedback loop that provides emotional context of a particular sensory input.[16]
While the dorsal hippocampus is involved in spatial memory formation, the left hippocampus is a
participant in the recall of these spatial memories. Eichenbaum[17] and his team found, when studying
the hippocampal lesions in rats, that the left hippocampus is "critical for effectively combining the
'what', 'when', and 'where' qualities of each experience to compose the retrieved memory". This
makes the left hippocampus a key component in the retrieval of spatial memory. However,
Spreng[18] found that the left hippocampus is a general concentrated region for binding together bits
and pieces of memory composed not only by the hippocampus, but also by other areas of the brain
to be recalled at a later time. Eichenbaum's research in 2007 also demonstrates that the
parahippocampal area of the hippocampus is another specialized region for the retrieval of
memories just like the left hippocampus.[citation needed]
Learning[edit]
The hippocampus, over the decades, has also been found to have a huge impact in learning. Curlik
and Shors[19] examined the effects of neurogenesis in the hippocampus and its effects on learning.
This researcher and his team employed many different types of mental and physical training on their
subjects, and found that the hippocampus is highly responsive to these latter tasks. Thus, they
discovered an upsurge of new neurons and neural circuits in the hippocampus as a result of the
training, causing an overall improvement in the learning of the task. This neurogenesis contributes to
the creation of adult-born granules cells (GC), cells also described by Eichenbaum[17] in his own
research on neurogenesis and its contributions to learning. The creation of these cells exhibited
"enhanced excitability" in the dentate gyrus (DG) of the dorsal hippocampus, impacting the
hippocampus and its contribution to the learning process.[17]
Hippocampus damage[edit]
Damage related to the hippocampal region of the brain has reported vast effects on overall cognitive
functioning, particularly memory such as spatial memory. As previously mentioned, spatial memory
is a cognitive function greatly intertwined with the hippocampus. While damage to the hippocampus
may be a result of a brain injury or other injuries of that sort, researchers particularly investigated the
effects that high emotional arousal and certain types of drugs had on the recall ability in this specific
memory type. In particular, in a study performed by Parkard,[20] rats were given the task of correctly
making their way through a maze. In the first condition, rats were stressed by shock or restraint
which caused a high emotional arousal. When completing the maze task, these rats had an impaired
effect on their hippocampal-dependent memory when compared to the control group. Then, in a
second condition, a group of rats were injected with anxiogenic drugs. Like the former these results
reported similar outcomes, in that hippocampal-memory was also impaired. Studies such as these
reinforce the impact that the hippocampus has on memory processing, in particular the recall
function of spatial memory. Furthermore, impairment to the hippocampus can occur from prolonged
exposure to stress hormones such as glucocorticoids (GCs), which target the hippocampus and
cause disruption in explicit memory.[21]
In an attempt to curtail life-threatening epileptic seizures, 27-year-old Henry Gustav
Molaison underwent bilateral removal of almost all of his hippocampus in 1953. Over the course of
fifty years he participated in thousands of tests and research projects that provided specific
information on exactly what he had lost. Semantic and episodic events faded within minutes, having
never reached his long-term memory, yet emotions, unconnected from the details of causation, were
often retained. Dr. Suzanne Corkin, who worked with him for 46 years until his death, described the
contribution of this tragic "experiment" in her 2013 book.[22]

Amygdala[edit]
Main article: Amygdala

Episodic-autobiographical memory (EAM) networks[edit]

Another integrative part of the limbic system, the amygdala, which is the deepest part of the limbic
system, is involved in many cognitive processes and is largely considered the most primordial and
vital part of the limbic system. Like the hippocampus, processes in the amygdala seem to impact
memory; however, it is not spatial memory as in the hippocampus but the semantic division of
episodic-autobiographical memory (EAM) networks. Markowitsch's[23] amygdala research shows it
encodes, stores, and retrieves EAM memories. To delve deeper into these types of processes by the
amygdala, Markowitsch[23] and his team provided extensive evidence through investigations that the
"amygdala's main function is to charge cues so that mnemonic events of a specific emotional
significance can be successfully searched within the appropriate neural nets and re-activated."
These cues for emotional events created by the amygdala encompass the EAM networks previously
mentioned.
Attentional and emotional processes[edit]
Besides memory, the amygdala also seems to be an important brain region involved in attentional
and emotional processes. First, to define attention in cognitive terms, attention is the ability to focus
on some stimuli while ignoring others. Thus, the amygdala seems to be an important structure in this
ability.
Foremost, however, this structure was historically thought to be linked to fear, allowing the individual
to take action in response to that fear. However, as time has gone by, researchers such as Pessoa,
[24]
 generalized this concept with help from evidence of EEG recordings, and concluded that the
amygdala helps an organism to define a stimulus and therefore respond accordingly. However,
when the amygdala was initially thought to be linked to fear, this gave way for research in the
amygdala for emotional processes. Kheirbek[15] demonstrated research that the amygdala is involved
in emotional processes, in particular the ventral hippocampus. He described the ventral
hippocampus as having a role in neurogenesis and the creation of adult-born granule cells (GC).
These cells not only were a crucial part of neurogenesis and the strengthening of spatial memory
and learning in the hippocampus but also appear to be an essential component to the function of the
amygdala. A deficit of these cells, as Pessoa (2009) predicted in his studies, would result in low
emotional functioning, leading to high retention rate of mental diseases, such as anxiety disorders.
[citation needed]

Social processing[edit]
Social processing, specifically the evaluation of faces in social processing, is an area of cognition
specific to the amygdala. In a study done by Todorov,[25] fMRI tasks were performed with participants
to evaluate whether the amygdala was involved in the general evaluation of faces. After the study,
Todorov concluded from his fMRI results that the amygdala did indeed play a key role in the general
evaluation of faces. However, in a study performed by researchers Koscik[26] and his team, the trait of
trustworthiness was particularly examined in the evaluation of faces. Koscik and his team
demonstrated that the amygdala was involved in evaluating the trustworthiness of an individual.
They investigated how brain damage to the amygdala played a role in trustworthiness, and found
that individuals with damaged amygdalas tended to confuse trust and betrayal, and thus placed trust
in those having done them wrong. Furthermore, Rule,[27] along with his colleagues, expanded on the
idea of the amygdala in its critique of trustworthiness in others by performing a study in 2009 in
which he examined the amygdala's role in evaluating general first impressions and relating them to
real-world outcomes. Their study involved first impressions of CEOs. Rule demonstrated that while
the amygdala did play a role in the evaluation of trustworthiness, as observed by Koscik in his own
research two years later in 2011, the amygdala also played a generalized role in the overall
evaluation of first impression of faces. This latter conclusion, along with Todorov's study on the
amygdala's role in general evaluations of faces and Koscik's research on trustworthiness and the
amygdala, further solidified evidence that the amygdala plays a role in overall social processing.
Klüver–Bucy syndrome[edit]
Main article: Klüver–Bucy syndrome

Based on experiments done on monkeys, the destruction of the temporal cortex almost always led to
damage of the amygdala. This damage done to the amygdala led the physiologists Kluver and Bucy
to pinpoint major changes in the behavior of the monkeys. The monkeys demonstrated the following
changes:

1. The monkeys were not afraid of anything.

2. The monkeys had extreme curiosity about everything.
3. The monkeys forgot rapidly.
4. The monkeys had a tendency to place everything in its mouth.
5. The monkeys often had a sexual drive so strong that they attempted to copulate with
immature animals, animals of the same sex, or even animals of a different species.
This set of behavioral change came to be known as the Klüver–Bucy syndrome.

Evolutionary claims[edit]
Paul D. MacLean, as part of his triune brain theory (which is now considered outdated[citation needed][28][29]),
hypothesized that the limbic system is older than other parts of the forebrain, and that it developed to
manage circuitry attributed to the fight or flight first identified by Hans Selye[30] in his report of the
General Adaptation Syndrome in 1936. It may be considered a part of survival adaptation in reptiles
as well as mammals (including humans). MacLean postulated that the human brain has evolved
three components, that evolved successively, with more recent components developing at the
top/front. These components are, respectively:

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.
2. The paleopallium or intermediate ("old mammalian") brain, comprising the structures of the
limbic system.
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, as a consequence, the human species. Similar
development of the neocortex in mammalian species unrelated to humans and primates has
also occurred, for example in cetaceans and elephants; thus the designation of "superior
mammals" is not an evolutionary one, as it has occurred independently in different species.
[dubious – discuss]
 The evolution of higher degrees of intelligence is an example of convergent
evolution, and is also seen in non-mammals such as birds.[citation needed]
According to Maclean, each of the components, although connected with the others, retained "their
peculiar types of intelligence, subjectivity, sense of time and space, memory, mobility and other less
specific functions".
However, while the categorization into structures is reasonable, the recent studies of the limbic
system of tetrapods, both living and extinct, have challenged several aspects of this hypothesis,
notably the accuracy of the terms "reptilian" and "old mammalian". The common ancestors of reptiles
and mammals had a well-developed limbic system in which the basic subdivisions and connections
of the amygdalar nuclei were established.[31] Further, birds, which evolved from the dinosaurs, which
in turn evolved separately but around the same time as the mammals, have a well-developed limbic
system. While the anatomic structures of the limbic system are different in birds and mammals, there
are functional equivalents.[citation needed]