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Neuroanatomy of episodic and semantic memory in humans: A brief review of

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Neuroanatomy of episodic and semantic
memory in humans: A brief review of
neuroimaging studies
Haydée G. García-Lázaro, Rocio Ramirez‑Carmona, Ruben Lara‑Romero1, Ernesto Roldan‑Valadez
Magnetic Resonance Unit, Medica Sur Clinic & Foundation, Mexico City, 1Division of Medicine and Health Sciences, University of
Sonora, Hermosillo City, Mexico
Address for correspondence:
Dr. Ernesto Roldan-Valadez,
Coordination of Research and
Innovation, Magnetic Resonance Unit,
Medica Sur Clinic and Foundation.
Puente de Piedra 150. Col. Toriello
Guerra. Deleg. Tlalpan, CP 14050,
Mexico City, Mexico.
E‑mail: ernest.roldan@usa.net
Received : 24‑11‑2012
Review completed : 25‑11‑2012
Accepted : 30‑11‑2012
Introduction
Memory is an essential function for the survival of
individuals and their species because the majority of
cognitive abilities depend on the successful storage of
information.[1] Memory has been defined as the process
of encoding, storing, consolidating, and retrieving
information. It is an emergent process in our brain,
resulting from complex interactions between the
biochemistry of neurons and their electrical activity in
specific anatomical structures.[2]
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Neurology India | Nov-Dec 2012 | Vol 60 | Issue 6
Topic of the issue: Review Article
Abstract
One of the most basic functions in every individual and species is memory. Memory is
the process by which information is saved as knowledge and retained for further use as
needed. Learning is a neurobiological phenomenon by which we acquire certain information
from the outside world and is a precursor to memory. Memory consists of the capacity to
encode, store, consolidate, and retrieve information. Recently, memory has been defined
as a network of connections whose function is primarily to facilitate the long‑lasting
persistence of learned environmental cues. In this review, we present a brief description
of the current classifications of memory networks with a focus on episodic memory
and its anatomical substrate. We also present a brief review of the anatomical basis of
memory systems and the most commonly used neuroimaging methods to assess memory,
illustrated with magnetic resonance imaging images depicting the hippocampus, temporal
lobe, and hippocampal formation, which are the main brain structures participating in
memory networks.
Key words: Hippocampus, magnetic resonance imaging, memory, neuroanatomy,
parahippocampal gyrus
Episodic memory (EM) stores information regarding
events and their context, such as what the event was,
and where and when it occurred.[3] It is important for
clinicians to know the structures involved in this kind
of memory because EM allows individuals remember
who they are, their ideas, thoughts, and feelings. Without
EM, we cannot learn from the past. It also gives us our
identity and individuality, and is an elementary part
of our consciousness. EM declines over the lifespan.
The elderly take more time to store information, and
their recall of recent events is less accurate than that of
young adults. Alterations to the structures underpinning
EM have been implicated in various disorders such as
Alzheimer’s disease,[4] epilepsy,[5] schizophrenia,[6] and
depression.[7]
Studies in cognitive neuroscience have demonstrated
that memory is a dynamic property of the brain as
a whole, rather than being localized to any single
region. [8‑12] New advances in magnetic resonance
613
 Garcia‑Lazaro, et al.: Neuroanatomy of episodic memory in humans
imaging (MRI) have allowed to obtain morphologic
and functional evidence of the different brain structures
involved in memory networks, which form interconnects
across the entire brain.[13] Because memory depends on
several brain systems working in concert across many
levels of neural organization, modern neuroscientists
can innovatively use MRI to determine the architecture
of memory systems.
We present a brief review of the current concepts and
classifications of EM and also show some selected
gray‑matter MR images from the hippocampus,
thalamus, amygdala, temporal lobe, hippocampal
formation, mammillary bodies, and cerebellum, as they
participate in EM.
Definitions
Memory is an important part of daily life, and allows us
to effectively interact with the environment and other
people. Memory allows for the retention of experiences,
and it minimizes risks and facilitates the optimal use
of the current environment. Memory consists of the
encoding, storage, consolidation, and retrieval of
information.[3,14,15]
Encoding refers to the initial registration or acquisition
of information. It involves the capture of information by
sensory systems and its conversion by neuronal coding
for use beyond simple perception.[16] The nature of
encoding may differ considerably depending on different
memory demands.
Storage is the creation of a relatively stable memory
trace or record of knowledge in the brain.[13] Such traces
require neuronal networks that can engage in neuronal
coding, which is the substrate of information storage, and
is evoked when we remember specific information.[17‑19]
Consolidation refers to slow physical processes, which
continue after perception, that enable temporary changes
in activity and synaptic strength to become long‑lasting,
and the later reactivation of neural activity to allow
for the induction of long‑term synaptic changes. This
occurs both in the region where the representation
was initially formed (e.g., the hippocampus) and in
additional regions where the representation was initially
weaker (e.g., neocortical structures), but which receive
spreading neural activity from the hippocampus.[20]
Retrieval refers to accessing stored information.[21] It
requires the reactivation of knowledge and is closely
related to encoding. Any successful act of retrieval
requires initial encoding and the persistence of
information in the nervous system.[22] Because these
processes require the active connection of several
brain structures, memory is not dependent on a
614
single anatomic structure to which we can assign
responsibility for an isolated memory process. Rather,
there are multiple memory systems that shape a widely
distributed network of cortical and subcortical brain
regions with specific neural behavior.
In the last few years, memory has been described as
a network of connections whose function is primarily
to facilitate the long‑lasting persistence of learned
environmental cues.[22] Memory systems are formed
by organizing elementary structures, which consist
of a neural substrate and its behavioral and cognitive
correlates. Some of these structures are engaged in
other memory systems, and they have emerged at
different stages of evolution and at different stages in
the development of organisms.[23]
Classifications of memory systems
Memory has been classified into many subtypes
depending on its persistence, the contents of its stored
material, and the presence or not of consciousness during
learning and memory.[1]
In accordance with its persistence, memory was first
separated into three sequential major systems, the
sensory, short‑term, and long‑term memory systems.[24]
Sensory memory allows for the recording of sensations
and their storage in cortical structures.[25] Short‑term
memory was initially proposed as a precursor to
long‑term memory. However, it was later noted that
not all information stored in short‑term memory passes
into long‑term memory. In fact, one kind of short‑term
memory refers to information used exclusively for
executing or developing other complex cognitive
processes.[26‑32]
Long‑term memory has been divided by Tulving[15] based
on whether the material is stored in EM or semantic
memory. EM is defined as recollections of previous
experiences from one’s personal past, especially if
focused on events that constitute our autobiographic
memory. Semantic memory refers to general knowledge
of facts and concepts regarding the world. This
knowledge is not located in a specific time or place.[3]
Based on the contents of their operating characteristics,
the kinds of information they process, and the purpose
they serve, all memory systems have been classified as
declarative and non‑declarative[1] [Figure 1]. Declarative
memory refers to memories that can be consciously
recalled, such as facts and events. It is divided into episodic
and semantic memory. In contrast, non‑declarative memory
does not afford awareness of any memory content but
does require consciousness, and it includes procedural
memory, priming, simple classical conditioning, and
non‑associative learning.[1]
Neurology India | Nov-Dec 2012 | Vol 60 | Issue 6
 Garcia‑Lazaro, et al.: Neuroanatomy of episodic memory in humans
Figure 1: Functional classification of memory systems, modified from
Squire[42] and van Strien[2]
Neuroimaging methods
The quest to obtain an understanding of the functional
organization of the normal human brain, using
techniques to assess changes in brain circulation, has
occupied neuroscientists for more than a century.
William James, in 1890, studied changes in brain flow
during mental activities. He based his work on the
writings of Angelo Mosso, who wrote in 1881 that brain
circulation changed selectively with neuron activity.[33] In
1950, Kety et al. provided the first glimpse of quantitative
changes in blood flow in the brain related directly to
brain function.[34]
Nowadays, cognitive neuroscience, a growing area in
neuroscience, combines the experimental strategies
of cognitive psychology with various neuroimaging
techniques to examine how brain functions support
mental activities. [35] The neuroimaging techniques
used include positron emission tomography (PET) and
magnetic resonance imaging (MRI).[36,37]
These neuroimaging methods provide a means of
measuring local changes in brain activity. PET uses
radiolabeled tracers to visualize blood flow changes
related to neural activity.[38] MRI can measure brain
activity indirectly by taking advantage of a fortuitous
physiologic property, which is that when a region of
the brain increases activity, both blood flow and the
oxygen content of the blood in that region increase.[39]
This allows for the indirect visualization of neural
activity through changes in oxygen content of the
blood. MRI can also provide quantitative analyses of
brain structure, so current quantitative MR reports
may include measurements of segmented volumes
from gray matter (GM), white matter (WM), and
cerebrospinal fluid (CSF), or even specific gyri.[40] Both
functional and structural MRI have the advantages
of being safe, providing excellent spatial resolution,
Neurology India | Nov-Dec 2012 | Vol 60 | Issue 6
Figure 2: Classification of memory systems based on their anatomy,
modified from Squire[42] and van Strien[2]
and not requiring the application of any intravenous
contrast agent.[37]
Anatomical basis of memory systems
In the last few decades, memory has come to be seen
as networks of interconnected cortical neurons, formed
by associations that contain our experiences in their
connectional structures.[12] Likewise, we now believe
that memory networks overlap and interact profusely
with one another such that a cellular assembly can be
engaged in many memory networks. These networks
are integrated by specific anatomical structures such
as the hippocampus, cerebellum, amygdala, frontal
lobes, temporal lobes, entorhinal cortex, and basal
ganglia[41] [Figure 2].
Brain structures associated with EM
Because EM deals with where information was acquired,
this subtype of declarative memory depends on the
integrity of the medial temporal lobe. The hippocampus
has a critical role in EM because it contains place cells,
head‑direction cells, and grid cells, which are involved
in the representation and recollection of spatial locations.
The hippocampus is surrounded and connected with the
entorhinal, parahippocampal, and perirhinal cortices,
and also with several subcortical and cortical structures.
The left prefrontal cortex is relatively highly involved in
encoding the information, whereas the right prefrontal
cortex is more involved in the EM retrieval.[23] The
hippocampal regions including the corpus of Ammon,
dentate gyrus, and subicular complex, and the adjacent
perirhinal, entorhinal, and parahippocampal cortices are
essential for declarative memory[42] [Figure 3].
Brain structures associated with semantic memory
Semantic memory involves the semantic definition of
objects. It uses shape, color, size, function, and motion
615
 Garcia‑Lazaro, et al.: Neuroanatomy of episodic memory in humans
a b
c d
Figure 3: Anatomical structures involved in episodic memory: (a) gyri
of the temporal lobe; (b and c) hippocampal formation and entorhinal
cortex; (d) frontal lobe
information. A consensus has not yet been achieved
regarding where this information is represented in the
brain. Some authors have proposed that this information
is stored in perceptual and motor systems, and was active
when we first learned about an object.[43] In support of
this argument, the occipital cortex is the beginning of
semantic processing, which continues in the left temporal
lobe [Figure 4]. The left inferior frontal cortex is relevant
to word selection and retrieval.[44] The fusiform gyrus,
located at the ventral surface of the temporal lobes, is
highly active during naming and reading words.[44] In
the Squire model, the fundamental memory structures
are the hippocampus, neocortex, amygdala, cerebellum,
and basal ganglia.[42]
Conclusions
The understanding of memory systems requires both
isolated diagrams of brain structures and knowledge of
the dynamic interconnections between these systems.
Anatomical and physiological assessments of brain
structures can be achieved with imaging techniques
such as PET and MRI. Knowledge of the relationships
between memory systems is not exclusively held by
neuroscientists doing research in humans or animals.
This knowledge is also available to clinicians currently
involved in the study of brain pathologies affecting
616
a b
c d
Figure 4: Brain structures involved in semantic memory:
(a, b and c) frontal, temporal, and occipital lobes (neocortex);
(d) proposed semantic memory network within the temporal
lobe, modified from Patterson et al.[44]
memory, such as neurologists, psychiatrists, geriatrists,
neurosurgeons, and psychologists. Although it is still
an evolving field, we believe that the evaluation of basic
systems requires easy and straightforward methods. So,
any clinician trained in these techniques could rapidly
obtain reliable and quantitative information that is useful
in clinical settings.
Acknowledgments
This study was supported in part by Medica Sur Clinic and
Foundation.
Haydée G. García-Lázaro was scholarship recipient of the
Institute of Science and Technology of Mexico City (ICyTDF
number EJE11-86) and worked at the MRI unit of Medica Sur
Clinic & Foundation from July 2011 to June 2012.
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How to cite this article: García-Lázaro HG, Ramirez-Carmona R,
Lara-Romero R, Roldan-Valadez E. Neuroanatomy of episodic and
semantic memory in humans: A brief review of neuroimaging studies.
Neurol India 2012;60:613-7.
Source of Support: Nill, Conflict of Interest: None declared.
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