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Anatomy of learning and

memory

Learning

is acquiring new or
modifying existing
knowledge, behaviors,
skills, valus, or preferences
and may involve
synthesizing different
types of information.
The ability to learn is
possessed by humans,
animals and some
machines.
Progress over time tends
to follow learning curves.

Human learning may occur as


part of education, personal
development, school or
training.
It may be goal-oriented and
may be aided by motivation.
The study of how learning
occurs is part of
neuropsychology, educational
psychology, learning theory,
and pedagogy.
Learning may occur as a
result of habituation or
classical conditioning, seen in
many animal species, or as a
result of more complex
activities such as play, seen
only in relatively intelligent
animals

Learning domains
Learning may occur
consciously or without
conscious awareness.
There is evidence for
human behavioral
learning prenatally, in
which habituation has
been observed as early
as 32 weeks into
gestation, indicating that
the central nervous
system is sufficiently
developed and primed
for learning and memory
to occur very early on in
development

Benjamin Bloom has


suggested three
domains of learning:
Cognitive To recall,
calculate, discuss,
analyze, problem solve,
etc.
Psychomotor To dance,
swim, ski, dive, drive a
car, ride a bike, etc.
Affective To like
something or someone,
love, appreciate, fear,
hate, worship, etc.

Memory: encode, store,


recall

has the ability to encode,


store and recall information.
Memories give an organism
the capability to learn and
adapt from previous
experiences as well as build
relationships.
Encoding allows the
perceived item of use or
interest to be converted
into a construct that can be
stored within the brain and
recalled later from short
term or long term memory.
Working memory stores
information for immediate
use or manipulation

Types:
Visual, acoustic, and
semantic encodings are
the most intensively used.
Other encodings are also
used.

encoding

Visual encoding
Visual encoding is the process of
encoding images and visual sensory
information.
Visual sensory information is
temporarily stored within our iconic
memory and working memory
before being encoded into
permanent long-term storage.
Baddeleys model of working
memory states that visual
information is stored in the visuospatial sketchpad.
The amygdala is a complex
structure that has an important role
in visual encoding. It accepts visual
input in addition to input from other
systems and encodes the positive or
negative values of conditioned
stimuli.

Acoustic encoding
Acoustic encoding is the processing
and encoding of sound, words, and all
other auditory input for storage and
later retrieval.

According to Baddeley, processing of


auditory information is aided by the
concept of the phonological loop,
which allows input within our echoic
memory to be sub vocally rehearsed
in order to facilitate remembering.

Studies indicate that lexical, semantic


and phonological factors interact in
verbal working memory. The
phonological similarity effect (PSE), is
modified by word concreteness.
This emphasizes that verbal working
memory performance cannot
exclusively be attributed to
phonological or acoustic
representation but also includes an
interaction of linguistic
representation.
What remains to be seen is whether
linguistic representation is expressed
at the time of recall or whether they
participate in a more fundamental
role in encoding and preservation.

encoding
Other senses
Tactile encoding is the
processing and encoding of
how something feels,
normally through touch.
Neurons in the primary
somatosensory cortex (S1)
react to vibrotactile stimuli
by activating in
synchronisation with each
series of vibrations.[6] Odors
and tastes may also lead to
encode.
In general encoding for
short-term storage (STS) in
the brain relies primarily on
acoustic rather than
semantic encoding

Semantic encoding
Semantic encoding is the
processing and encoding of
sensory input that has
particular meaning or can
be applied to a context.
Various strategies can be
applied such as chunking
and mnemonics to aid in
encoding, and in some
cases, allow deep
processing, and optimizing
retrieval.

Brodmanns areas 45, 46, and


47 (the left inferior prefrontal
cortex or LIPC) showed
significantly more activation
during semantic encoding
conditions compared to
nonsemantic encoding
conditions regardless of the
difficulty of the nonsemantic
encoding task presented.
The same area showing
increased activation during
initial semantic encoding will
also display decreasing
activation with repetitive
semantic encoding of the
same words

Long term potentiation


Encoding is a biological event
that begins with perception
All perceived and striking
sensations travel to your brains
hippocampus where all these
sensations are combined into
one single experience.
The hippocampus is responsible
for analyzing these inputs and
ultimately deciding if they will be
committed to your long term
memory; these various threads
of information are stored in
various parts of the brain.
However, the exact way in which
these pieces are identified and
recalled later remains unknown.

Encoding

is achieved using a combination of


chemicals and electricity.
Neurotransmitters are released
when an electrical pulse crosses
the synapse which serves as a
connection from nerve cells to
other cells.
The dendrites receive these
impulses with their feathery
extensions.
A phenomenon called Long Term
Potentiation allows a synapse to
increase strength with increasing
numbers of transmitted signals
between the two neurons.
These cells also organise
themselves into groups specializing
in different kinds of information
processing. Thus, with new
experiences your brain creates
more connections and may
rewire.

The brain organizes and


reorganizes itself in response
to your experiences, creating
new memories prompted by
experience, education, or
training.
Therefore the use of a brain
reflects how it is organised.
This ability to re-organize is
especially important if ever a
part of your brain becomes
damaged.
Scientists are unsure of
whether the stimuli of what
we do not recall are filtered
out at the sensory phase or if
they are filtered out after the
brain examines their
significance.

Positron emission tomography (PET)


demonstrates a consistent
functional anatomical blueprint of
hippocampal activation during
episodic encoding and retrival.
Activation in the hippocampal region
associated with episodic memory
encoding has been shown to occur
in the rostral portion of the region
whereas activation associated with
episodic memory retrieval occurs in
the caudal portions.

This is referred to as the


Hippocampal Encoding/Retrieval
model or HIPER model.

One study used PET to measure


cerebral blood flow during encoding
and recognition of faces in both
young and older participants. Young
people displayed increased cerebral
blood flow in the right hippocampus
and the left prefrontal and temporal
cortices during encoding and in the
right prefrontal and parietal cortex
during recognition.

Mapping
activity
Elderly people showed no significant
activation in areas activated in young
people during encoding, however they
did show right prefrontal activation
during recognition.
Thus it may be concluded that as we
grow old, failing memories may be the
consequence of a failure to adequately
encode stimuli as demonstrated in the
lack of cortical and hippocampal
activation during the encoding
process.
Recent findings in studies focusing on
patients with posttraumatic stress
disorder demonstrate that amino acid
transmitters, glutamate and GABA, are
intimately implicated in the process of
factual memory registration, and
suggest that amine neurotransmitters,
norepinephrine and serotonin, are
involved in encoding emotional
memory.

Molecular perspective
The process of encoding is
not yet well understood,
however key advances
have shed light on the
nature of these
mechanisms.
Encoding begins with any
novel situation, as the
brain will interact and draw
conclusions from the
results of this interaction.
These learning
experiences have been
known to trigger a cascade
of molecular events
leading to the formation of
memories.

these changes include


the modification of neural
synapses, modification of
proteins, creation of new
synapses, activation of
gene expression and new
protein synthesis.
However, encoding can
occur on different levels.
The first step is shortterm memory formation,
followed by the
conversion to a long-term
memory, and then a
long-term memory
consolidation process.

Synaptic plasticity

Synaptic plasticity is the


ability of the brain to
strengthen, weaken, destroy
and create neural synapses
and is the basis for learning.

These molecular distinctions


will identify and indicate the
strength of each neural
connection.

The effect of a learning


experience depends on the
content of such an experience.
Reactions that are favoured
will be reinforced and those
that are deemed unfavourable
will be weakened.

This shows that the synaptic


modifications that occur can
operate either way, in order
to be able to make changes
over time depending on the
current situation of the
organism.
In the short term, synaptic
changes may include the
strengthening or weakening of
a connection by modifying the
preexisting proteins leading to
a modification in synapse
connection strength.
In the long term, entirely new
connections may form or the
number of synapses at a
connection may be increased,
or reduced.

The encoding
process

A significant short-term biochemical


change is the covalent modification
of pre-existing proteins in order to
modify synaptic connections that
are already active.
This allows data to be conveyed in
the short term, without
consolidating anything for
permanent storage.
From here a memory or an
association may be chosen to
become a long-term memory, or
forgotten as the synaptic
connections eventually weaken.
The switch from short to long-term
is the same concerning both implicit
memory and explicit memory
This process is regulated by a
number of inhibitory constraints,
primarily the balance between
protein phosphorylation and
dephosphorylation

Finally, long term changes occur that


allow consolidation of the target
memory.
These changes include new protein
synthesis, the formation of new
synaptic connections and finally the
activation of gene expression in
accordance with the new neural
configuration.
The encoding process has been
found to be partially mediated by
serotonergic interneurons,
specifically in regard to sensitization
as blocking these interneurons
prevented sensitization entirely.
However, the ultimate consequences
of these discoveries have yet to be
identified. Furthermore, the learning
process has been known to recruit a
variety of modulatory transmitters in
order to create and consolidate
memories.
These transmitters cause the nucleus
to initiate processes required for
neuronal growth and long term
memory, mark specific synapses for
the capture of long-term processes,
regulate local protein synthesis and
even appear to mediate attentional
processes required for the formation
and recall of memories.

Storage

in human memory is one of three


core process of memory, along with
Recall and Encoding.
It refers to the retention of
information, which has been
achieved through the encoding
process, in brain for prolonged
period of time until it is accessed by
the recall process.
Modern memory psychology
differentiates the two distinct type
of memory storage; short-term
memory and long-term memory.
In addition, different memory
models have suggested variations
of existing short-term and long-term
memory to account for different
ways of storing memory.

Models of Memory Storage


Varieties of different memory
models have been proposed to
account for different type of recall
process, including the cued recall,
free recall, and serial recall.
In order to explain the recall
process, however, the memory
model must identify how an
encoded memory can reside in the
memory storage for prolonged
period of time until the memory is
accessed again, during the recall
process. Not all models, however,
use the terminology of Short-Term
and Long-Term Memory to explain
the memory storage; the Dual-Store
theory and refined version of
Atkinson-Shiffrin Model of Memory
(Atkinson 1968) uses both ShortTerm and Long-Term memory
storage, but others do not.

Recall
in memory refers to
the retrieval of events
or information from
the past. Along with
encoding and storage,
it is one of the three
core processes of
memory.

There are three main


types of recall: free
recall, cued recall and
serial recall.
Psychologists test these
forms of recall as a way
to study the memory
processes of humans
and animals.
Two main theories of
the process of recall are
the Two-Stage Theory
and the theory of
Encoding Specificity.

Two-stage theory
The two-stage theory states
that the process of recall begins
with a search and retrieval
process, and then a decision or
recognition process where the
correct information is chosen
from what has been retrieved.
In this theory, recognition only
involves the latter of these two
stages, or processes, and this is
thought to account for the
superiority of the recognition
process over recall.
Recognition only involves one
process in which error or failure
may occur, while recall involves
two.
However, recall has been found
to be superior to recognition in
some cases, such as a failure to
recognize words that can later
be recalled.

recall

Encoding specificity
The theory of encoding
specificity finds similarities
between the process of
recognition and that of recall.
The encoding specificity
principle states that memory
utilizes information from the
memory trace, or the situation
in which it was learned, and
from the environment in which
it is retrieved.
Encoding specificity helps to
take into account context cues
because of its focus on the
retrieval environment, and it
also accounts for the fact
recognition may not always be
superior to recall

Brain Structures involve in


memory
The neuroanatomy
of memory
encompasses a wide
variety of
anatomical
structures in the
brain.

Subcortical
structures
Hippocampus
Cerebellum
Amygdala
Basal ganglia
Cortical structures
Frontal lobe
Temporal lobe
Parietal lobe
Occipital lobe

hippocampus
The hippocampus is
located in the medial
temporal lobe of the
brain
In this lateral view of
the human brain, the
frontal lobe is at left,
the occipital lobe at
right, and the temporal
and parietal lobes
have largely been
removed to reveal the
hippocampus
underneath.

Hippocampus
The hippocampus
The hippocampus is a
structure in the brain
that has been associated
with various memory
functions. It is part of the
limbic system, and lies
next to the medial
temporal lobe. It is made
up of two structures, the
Ammons Horn, and
the Dentate gyrus, each
containing different
types of cells

Cognitive maps
There is evidence that
the hippocampus
contains cognitive maps
in humans. In one study,
single-cell recordings
were taken from
electrodes implanted in
a rats hippocampus,
and it was found that
certain neurons
responded strongly only
when the rat was in
certain locations.

These cells are called place


cells, and collections of these
cells can be considered to be
mental maps. Individual place
cells do not only respond to
one unique area only however,
the patterns of activation of
these cells overlap to form
layered mental maps within
the hippocampus.
A good analogy is the
example of the same
television or computer screen
pixels being used to light up
any trillions of possible
combinations to produce
images, just as the place cells
can be used in any multiple
possible combinations to
represent mental map

The hippocampus right side


is more oriented towards
responding to spatial aspects,
whereas the left side is
associated with other context
information.
Also, there is evidence that
experience in building
extensive mental maps, such
as driving a city taxi for a long
time (since this requires a lot
of memorization of routes),
can increase the volume of
ones hippocampus.

Encoding
Damage to the hippocampus
and surrounding area can
cause anterograde amnesia,
the inability to form new
memories.
This implies that the
hippocampus is important not
only for storing cognitive
maps, but for encoding
memories

The hippocampus is also involved


in memory consolidation, the
slow process by which memories
are converted from short to long
term memory
This is supported by studies in
which lesions are applied to rat
hippocampi at different times
after learning.
The process of consolidation may
take up to a couple years.
It has also been found that it is
possible to form new semantic
memories without the
hippocampus, but not episodic
memories, which means that
explicit descriptions of actual
events (episodic) cannot be
learned, but some meaning and
knowledge is gained from
experiences (semantic).

hippocampus

The hippocampus

is a major component of the


brains of humans and other
vertebrates.
It belongs to the limbic system
and plays important roles in the
consolidation of information from
short-term memory to
long-term memory and spatial
navigation.
Like the cerebral cortex, with
which it is closely associated, it is
a paired structure, with mirrorimage halves in the left and right
sides of the brain.
In humans and other primates,
the hippocampus is located inside
the medial temporal lobe,
beneath the cortical surface.
It contains two main interlocking
parts: Ammon's horn and the
dentate gyrus.

In Alzheimer's disease, the


hippocampus is one of the
first regions of the brain to
suffer damage; memory
problems and disorientation
appear among the first
symptoms. Damage to the
hippocampus can also result
from oxygen starvation (
hypoxia), encephalitis, or
medial temporal lobe epilepsy
. People with extensive,
bilateral hippocampal damage
may experience
anterograde amnesiathe
inability to form or retain new
memories

Since different neuronal cell


types are neatly organized
into layers in the
hippocampus, it has
frequently been used as a
model system for studying
neurophysiology. The form of
neural plasticity known as
long-term potentiation (LTP)
was first discovered to occur
in the hippocampus and has
often been studied in this
structure. LTP is widely
believed to be one of the
main neural mechanisms by
which memory is stored in the
brain

The earliest description of the


ridge running along the floor
of the
temporal horn of the lateral ve
ntricle
comes from the Venetian
anatomist Julius Caesar Aranzi
(1587), who initially likened it
to a seahorse, using the Latin:
hippocampus (from Greek:
, "horse" and Greek:
, "sea monster") or
alternatively to a silkworm.

the German anatomist


Duvernoy (1729), the first to
illustrate the structure, also
wavered between "seahorse"
and "silkworm." "Ram's horn"
was proposed by the Danish
anatomist Jacob Winslw in
1732; and a decade later his
fellow Parisian, the surgeon de
Garengeot, used "cornu
Ammonis" - horn of (the
ancient Egyptian god) Amun

hippocampus

The hippocampus
as a whole has the
shape of a curved
tube, which has
been analogized
variously to a
seahorse, a ram's
horn (Cornu
Ammonis, hence the
subdivisions CA1
through CA4), or a
banana.[3

Today, the structure is


called the hippocampus
rather than
hippocampus major,
with pes hippocampi
often being regarded as
synonymous with De
Garengeot's "cornu
Ammonis",[2] a term
which survives in the
names of the four main
histological divisions of
the hippocampus: CA1,
CA2, CA3 and CA4

hippocampu
s
Over the years,
three main ideas of
hippocampal
function have
dominated the
literature: inhibition,
memory, and space.

The major line of thought relates


the hippocampus to memory.
Although it had historical
precursors, this idea derived its
main impetus from a famous
report by Scoville and
Brenda Milner[12] describing the
results of surgical destruction of
the hippocampus (in an attempt
to relieve epileptic seizures), in
a patient named Henry Gustav
Molaison,[13] known until his
death in 2008 as H.M. The
unexpected outcome of the
surgery was severe anterograde
and partial retrograde amnesia
The important theory of
hippocampal function relates
the hippocampus to space. The
spatial theory was originally
championed by O'Keefe and
Nadel, who were influenced by
E.C. Tolman's theories about "
cognitive maps" in humans and
animals

Role
hippocampu
s
that animals with

hippocampal damage tend to


be hyperactive; second, that
animals with hippocampal
damage often have difficulty
learning to inhibit responses
that they have previously
been taught, especially if the
response requires remaining
quiet as in a passive
avoidance test. Jeffrey Gray
developed this line of thought
into a full-fledged theory of
the role of the hippocampus in
anxiety.[10] The inhibition
theory is currently the least
popular of the three.[11]

Role in memory
Psychologists and
neuroscientists generally
agree that the
hippocampus has an
important role in the
formation of new memories
about experienced events (
episodic or
autobiographical memory).
Part of this role is
hippocampal involvement
in the detection of novel
events, places and stimuli.

Some researchers view


the hippocampus as
part of a larger
medial temporal lobe
memory system
responsible for general
declarative memory
(memories that can be
explicitly verbalized
these would include, for
example,
memory for facts in
addition to episodic
memory).

hippocampu
s
Location:medial to the
temporal horn of the
lateral ventriclein the
deep anterior temporal
lobe
Architecture:seahorse
shape structure
composeof the dentate
gyrus, CA regions, and
subiculum. It has 3
layer neuronal layers

Circuitry:
Internal
circuitry:entorhinal
cortex-dentate gyrusCA3-CA1-subiculumentorhinal cortex
Major external circuit:
Papez circuit:
hippocampus-fornixmammilary bodyanterior nucleus of
thalamus-cingulate
cortex-temporal
cortex-hippocampus

Inputs
Septal nuclei-fornix (cholinergic
input vital for memory and
degerates in alzheimer`s
disease
Multiple cortical areasentorhinal cortex-dentate gyrus
Inputs to entorhinal: sensory
assosiation
cortices,prefronatal,amygdala,o
lfactory
Inputs to subiculum:
amygdala,cingulate cortex
Inputs to hippocampus:raphe
nuclei (5HT),locus coeruleus
(NE)

Inputs outputs
Outputs
Subiculum-fornixmammillary body
CA1/CA3-fornixseptal
nuclei,nucleus
accumbens,hypot
halamus,cingulat
e cortex,frontal
lobe

basal ganglia (or basal


nuclei)

are a group of nuclei of


varied origin (mostly
telencephalic embryonal
origin, with some
diencephalic and
mesencephalic elements)
in the brains of
vertebrates that act as a
cohesive functional unit.
They are situated at the
base of the forebrain and
are strongly connected
with the cerebral cortex,
thalamus and other brain
areas.

The basal ganglia are


associated with a variety of
functions, including
voluntary motor control,
procedural learning relating
to routine behaviors or
"habits" such as bruxism,
eye movements, and
cognitive,emotional
functions.
Currently popular theories
implicate the basal ganglia
primarily in action selection
, that is, the decision of
which of several possible
behaviors to execute at a
given time.

location

Main
component
s

The main components of the


basal ganglia are the striatum
(also called neostriatum)
composed of caudate and
putamen, globus pallidus or
pallidum composed of globus
pallidus externa (GPe) and
globus pallidus interna (GPi),
substantia nigra composed of
both substantia nigra pars
compacta (SNc) and
substantia nigra pars

reticulata (SNr), and the


subthalamic nucleus (STN).

The largest component, the striatum,


receives input from many brain areas
but sends output only to other
components of the basal ganglia.
The pallidum receives its most
important input from the striatum
(either directly or indirectly), and
sends inhibitory output to a number
of motor-related areas, including the
part of the thalamus that projects to
the motor-related areas of the cortex.
One part of substantia nigra, the
reticulata (SNr), functions similarly to
the pallidum, and another part
(compacta or SNc) provides the
source of the neurotransmitter
dopamine's input to the striatum.
The subthalamic nucleus (STN)
receives input mainly from the
striatum and cortex, and projects to a
portion of the pallidum (interna
portion or GPi).
Each of these areas has a complex
internal anatomical and
neurochemical organization.

Role basal ganglia

The basal ganglia play a


central role in a number of
neurological conditions,
including several
movement disorders. The
most notable are
Parkinson's disease, which
involves degeneration of
the melanin-pigmented
dopamine-producing cells
in the substantia nigra pars
compacta (SNc), and
Huntington's disease,
which primarily involves
damage to the striatum.

Basal ganglia dysfunction is


also implicated in some other
disorders of behavior control
such as the
Tourette's syndrome, ballismus
(particularly hemibalismus),
obsessivecompulsive disorder
(OCD), and Wilson's disease
(Hepatolenticular
degeneration); except for
Wilson's disease and
hemiballismus, the
neuropathological mechanisms
underlying diseases of ganglia
such as Parkinsons' and
Huntington's are not very well
understood or are at best still
developing theories.[citation needed]

location

4 parts
In terms of anatomy, the basal ganglia are divided by
anatomists into four distinct structures, depending on
how superior or rostral they are (in other words
depending on how close to the top of the head they
are):
Two of them, the striatum and the pallidum, are
relatively large;
the other two, the substantia nigra and the
subthalamic nucleus, are smaller.
In the illustration to the right, two coronal sections of
the human brain show the location of the basal ganglia
components.
Of note, and not seen in this section, the subthalamic
nucleus and substantia nigra lie farther back (
posteriorly) in the brain than the striatum and pallidum.

the structures relevant to the


basal ganglia are shown in bold
Telencephalon: The
human cortex (both
hemispheres),
Caudate, Putamen,
Globus pallidus
(pallidum)
Diencephalon:
Thalamus,
hypothalamus,
subthalamus,
epithalamus,
subthalamic nuclei

Mesencephalon
(Midbrain),
Substantia nigra
pars compacta
(SNc), Substantia
nigra pars
reticulata (SNr)

Circuit connections
excitatory glutamatergic
pathways, inhibitory
GABAergic pathways, and
modulatory dopaminergic
pathways (Abbreviations: GPe:
globus pallidus external; GPi:
globus pallidus internal; STN:
subthalamic nucleus; SNc:
substantia nigra compacta;
SNr: substantia nigra
reticulata)
In order to understand the
complex circuitry of the basal
ganglia, one has to first
understand the important
participants in this circuit.
Parts of the basal ganglia are
in direct communication with
the thalamus and the cortex.
The cortex, thalamus, and the
basal ganglia are, therefore,
the three main participants in
the circuit created by the
basal ganglia.

circuit

At the top of the hierarchy lies


the cerebral cortex. The cortex
has many different areas with
different functions. One such
cortical area is called the
primary motor cortex (along the
pre-central gyrus). Specialized
neurons from the primary motor
cortex extend their axons all
the way to the striatum portion
of the basal ganglia.
These cortical neurons release
the neurotransmitter glutamate
, which is excitatory in nature.
Once excited by glutamate, the
cells in the striatum project in
two different directions giving
rise to two major pathways: The
"direct" and the "indirect"
pathways:

The following diagram


depicts the direct
pathway:
Cortex (stimulates)
Striatum (inhibits)
"SNr-GPi" complex
(less inhibition of
thalamus)
Thalamus
(stimulates) Cortex
(stimulates)
Muscles, etc.
(hyperkinetic state)

Cortex (stimulates)
Striatum (inhibits)
GPe (less inhibition of
STN) STN
(stimulates) "SNrGPi" complex
(inhibits) Thalamus
(is stimulating less)
Cortex (is stimulating
less) Muscles, etc.
(hypokinetic state)

Parkinson vs huntington
disease
Information about the functions of the basal ganglia
comes from anatomical studies, from physiological
studies carried out mainly in rats and monkeys, and
from the study of diseases that damage them.
The greatest source of insight into the functions of the
basal ganglia has come from the study of two
neurological disorders, Parkinson's disease and
Huntington's disease.
For both of these disorders, the nature of the neural
damage is well understood and can be correlated with
the resulting symptoms. Parkinson's disease involves
major loss of dopaminergic cells in the substantia nigra;
Huntington's disease involves massive loss of medium
spiny neurons in the striatum.

The symptoms of the two diseases are virtually


opposite: Parkinson's disease is characterized by
gradual loss of the ability to initiate movement, whereas
Huntington's disease is characterized by an inability to
prevent parts of the body from moving unintentionally.

It is noteworthy that, although both diseases have


cognitive symptoms, especially in their advanced
stages, the most salient symptoms relate to the ability
to initiate and control movement. Thus, both are
classified primarily as movement disorders.
A different movement disorder, called hemiballismus,
may result from damage restricted to the subthalamic
nucleus. Hemiballismus is characterized by violent and
uncontrollable flinging movements of the arms and legs.

The role in motivation of the "limbic" part of the basal


gangliathe nucleus accumbens (NA),
ventral pallidum, and ventral tegmental area (VTA)is
particularly well established. Thousands of
experimental studies combine to demonstrate that the
dopaminergic projection from the VTA to the NA plays
a central role in the brain's reward system.
Animals with stimulating electrodes implanted along
this pathway will bar-press very energetically if each
press is followed by a brief pulse of electrical current.
Numerous things that people find rewarding, including
addictive drugs, good-tasting food, and sex, have
been shown to elicit activation of the VTA dopamine
system. Damage to the NA or VTA can produce a state
of profound torpor.

Basal ganglia
Involved in initiation and modulation
of movement, receive input from
cerebral cortex, process it, and relay
back to cerebral cortex via thalamus
Consists: corpus striatum (striatum +
pallidum)(striatum:nucleus caudatus+
putamen) ,globus pallidus (pallidum,
+ putamen: nucleus lentiformis);
amygdala, claustrum

Nucleus caudatus
C shaped, is lateral to the lateral ventricle,
composes: head, body and tail
Head, bulges into the frontal horn of lateral
ventricle, connect to the lentiform nucleus;
body sweeps into a c shaped with the
lateral ventricle, capsula interna ;tail
located in the temporal lobe superior and
lateral to the temporal horn of the lateral
ventricle, connect to amigdaloid body
Degeneration lead : Huntington`s disesae

Putamen & globus pallidus


Lens shaped, lateral to globus
pallidus, anteriorly and inferiorly is
connected to nucleus caudatus
Corpus striatum: putamen + nucleus
caudatus
Globus pallidus- pale globe
Nucleus lentiformis= putamen +
globus pallidus

Substantia nigra
Located in the ventral midbrain, appears brown,
neuromelanin pigmentation
Pars compacta- has dopamine producing neuron,
project to striatum and facilitate movement
Pars reticularis-is an output nucleus
Degeneration causes: parkinson`s disease;
Chorea=lesion is above and medial subtantia
nigra; athetosis= it is above and lateral to it
Ballism, hemibalismus =subthalamic nucleus;
tremor =lesion between subtantia nigra and red
nucleus

Basal ganglia and motor


memory
The basal ganglia
The basal ganglia are a
group of nuclei which are
located in the medial
temporal lobe, above the
thalamus and connected to
the cerebral cortex.
Specifically, the basal
ganglia includes the
subthalamic nucleus,
substantia nigra, the
globus pallidus, the ventral
striatum and the dorsal
striatum, which consists of
the putamen and the
caudate nucleus

The basic functions of


these nuclei deal with
cognition, learning, and
motor control and
activities.
The basal ganglia are also
associated with learning,
memory, and unconscious
memory processes, such
as motor skills and implicit
memory

The caudate nucleus is


thought to assist in
learning and memory of
associations taught during
operant conditioning.
Specifically, research has
shown that this part of the
basal ganglia plays a role
in acquiring stimulusresponse habits, as well as
in solving sequence tasks

Damage to the basal ganglia


has been linked to
dysfunctional learning of
motor and perceptual-motor
skills.
Most disorders that are
associated with damage to
these areas of the brain
involve some type of motor
dysfunction, as well as
trouble with mental
switching between tasks in
working memory
Such symptoms are often
present in those who suffer
from dystonia, ,
athymhormic syndrome,
Fahr's syndrome,
Huntington's disease or
Parkinson's disease.
Huntington's and Parkinson's
disease involve both motor
deficits and cognitive
impairment

amygdala
location

Subdivision

amygdala
The amygdalae (/mdli/
; singular: amygdala; also
corpus amygdaloideum; Latin
, from Greek ,
amygdal, 'almond', 'tonsil',
listed in the Gray's Anatomy
as the nucleus amygdal)
[1] are almond-shaped groups
of nuclei located deep within
the medial temporal lobes of
the brain in complex
vertebrates, including
humans.
Shown in research to perform
a primary role in the
processing and memory of
emotional reactions, the
amygdalae are considered
part of the limbic system.

Connections
The amygdala sends
impulses to the
hypothalamus for activation
of the
sympathetic nervous system,
to the
thalamic reticular nucleus for
increased reflexes, to the
nuclei of the trigeminal nerve
and the facial nerve, and to
the ventral tegmental area,
locus coeruleus, and
laterodorsal tegmental nucleu
s
for activation of dopamine,
norepinephrine and
epinephrine.
Coronal section of brain
through intermediate mass of
third ventricle.

The cortical nucleus is


involved in the sense of
smell and pheromoneprocessing.
It receives input from the
olfactory bulb and
olfactory cortex.
The lateral amygdalae, which
send impulses to the rest of
the basolateral complexes
and to the centromedial
nuclei, receive input from the
sensory systems. The
centromedial nuclei are the
main outputs for the
basolateral complexes, and
are involved in emotional
arousal in rats and cats.

amygdala

Role amygdala

during fear conditioning, sensory stimuli reach the basolateral complexes of the
amygdalae, particularly the lateral nuclei, where they form associations with
memories of the stimuli. The association between stimuli and the aversive
events they predict may be mediated by long-term potentiation, a sustained
enhancement of signalling between affected neurons.
Memories of emotional experiences imprinted in reactions of synapses in the
lateral nuclei elicit fear behavior through connections with the central nucleus
of the amygdalae and the bed nuclei of the stria terminalis (BNST). The central
nuclei are involved in the genesis of many fear responses, including freezing
(immobility), tachycardia (rapid heartbeat), increased respiration, and stresshormone release. Damage to the amygdalae impairs both the acquisition and
expression of Pavlovian fear conditioning, a form of classical conditioning of
emotional responses.[3]
The amygdalae are also involved in appetitive (positive) conditioning. It seems
that distinct neurons respond to positive and negative stimuli, but there is no
clustering of these distinct neurons into clear anatomical nuclei.
However, lesions of the central nucleus in the amygdala have been shown to
reduce appetitive learning in rats. Lesions of the basolateral lesions do not
exhibit the same effect.
Research like this indicates that different nuclei within the amygdala have
different functions in appetitive conditioning

Amygdala
The amygdala
Located below the
hippocampus in the
medial temporal lobes
are two amygdalae
(singular "amygdala").
The amygdala are
associated with both
emotional learning and
memory, as it
responds strongly to
emotional stimuli,
especially fear.

These neurons assist in


encoding emotional
memories and
enhancing them.
This process results in
emotional events being
more deeply and
accurately encoded into
memory.
Lesions to the
amygdalae in monkeys
have been shown to
impair motivation, as
well as the processing
of emotions

Memory of fear
conditioning
Pavlovian conditioning
tests have shown the
active role of the
amygdala in fear
conditioning in rats.
Research involving
lesions to the basolateral
nucleus have shown a
strong association with
memories involving fear.
The central nucleus is
linked with the behavioral
responses that are
dependent on the
basolaterals reaction to
fear.

The central nucleus of


the amygdala is also
linked to emotions and
behaviors motivated by
food and sex

Memory consolidation
Emotional experiences and
events are somewhat fragile
and take a while to be
completely set into memory.
This slow process, referred
to as consolidation, allows
emotions to influence the
way the memory is stored.
The amygdala is involved in
memory consolidation,
which is the process of
transferring information that
is currently in working
memory into ones long-term
memory. This process is also
known as memory
modulation.

The amygdala works to


encode recent emotional
information into memory.
Memory research has shown
that the greater ones
emotional arousal level at
the time of the event, the
greater the chance that the
event will be remembered.
This may be due to the
amygdala enhancing the
emotional aspect of the
information during encoding,
causing the memory to be
processed at a deeper level
and therefore, more likely to
withstand forgetting.

amygdala
Almond shaped, located anterior to the hippocampus
in the anterior portion of medial temporal lobe
Inputs:major= temporal lobe, olfactory, limbic,
autonomic information from orbitofrontal, cingulate
gyrus, hypothalamus, tegmentum, brainstem: DA
(ventral tegmental area), NE(raphe nuclei),
5HT(locus coeruleus)
Outputs: major= hypothalamus, thalamus,
striatum,septal nuclei, hippocampus, multiple
cortical areas
Function: regulates emotional interpretation, relating
to fear and anger

Limbic system
A system of nuclear structures and tracts
found in a ring that encircles the thalamus
Multiple structures interact with each other
and with other cortical and subcortical
structure, function serves major regulator of
emotional control and memory encoding
Damage can lead aggression, apathy,
anterograde amnesia (inability to form new
memories)

Major limbic forebrain


structures
Hippocampus: memory encoding

Fornix:C shaped, extend hippocampus travels inferior to


corpus callosum-dives down to mamillary body
Amygdala
Stria terminalis, c shaped tract connecting amygdala to
the hypothalamus and basal forebrain
Habenula, small nuclear structure, rostral the pineal
gland, part of epithalamus
Septal nuclei, set of nuclear structurs rostral to anterior
commisura, as a pleasure centre of the brain; connection
with hippocampus, amygdala, and hypothalamus
Cingulate cortex:above corpus callosum, involve in
papez circuit, function in cortical regulation of autonomic
function, behavior, emotional modulation pain

Gyrus cinguli

Frontal lobe

The frontal lobes are located


at the front of each cerebral
hemisphere and positioned
anterior to the parietal lobes.
It is separated from the
parietal lobe by the primary
motor cortex, which controls
voluntary movements of
specific body parts associated
with the precentral gyrus.
The cortex here serves our
ability to plan the day,
organize work, type a letter,
pay attention to details and
control the movements of
your arms and legs.
It also contributed to your
personality and behaviour.

When considering the frontal


lobes in regards to memory, we
see that it is very important in
the coordination of information.

Therefore, the frontal lobes are


important in working memory,
For example, when you are
thinking about how to get to a
mall you have never been to
before, you combine various
bits of knowledge you already
have: the layout of the city the
mall is in, information from a
map, knowledge of traffic
patterns in that area and
conversations with your friends
about the location of the mall.
By actively using all of this
information, you can determine
the best route for you to take.
This action involves the
controlled use of information in
working memory, coordinated
by the frontal lobes.

The frontal lobe contains most of the


dopamine-sensitive neurons in the
cerebral cortex. The dopamine system is
associated with reward, attention, short-term
memory tasks, planning, and drive. Dopamine
tends to limit and select sensory information
arriving from the thalamus to the fore-brain. A
report from the National Institute of Mental
Health says a gene variant that reduces
dopamine activity in the prefrontal cortex is
related to poorer performance and inefficient
functioning of that brain region during working
memory tasks, and to slightly increased risk for
schizophrenia.

The executive functions of the frontal lobes involve the


ability to recognize future consequences resulting from
current actions, to choose between good and bad
actions (or better and best), override and suppress
unacceptable social responses, and determine
similarities and differences between things or events.
Therefore, it is involved in higher mental functions.
The frontal lobes also play an important part in
retaining longer term memories which are not taskbased. These are often memories associated with
emotions derived from input from the brain's
limbic system. The frontal lobe modifies those emotions
to generally fit socially acceptable norms.
Psychological tests that measure frontal lobe function
include finger tapping, Wisconsin Card Sorting Task,
and measures of verbal and figural fluency.[2]

The frontal lobes help a


person select out
memories that are most
relevant on a given
occasion. It can
coordinate various types
of information into a
coherent memory trace
For example, the
knowledge of the
information itself, as well
as knowing where
information came from
must be put together into
a single memory
representation; this is
called source monitoring.

Sometimes we experience
situations where
information becomes
separated, such as when
we recall something, but
cannot remember where
we remember it from; this
is referred to as a source
monitoring error
The frontal lobes are also
involved in the ability to
remember what we need
to do in the future; this is
called prospective
memory

Temporal lobe
The temporal lobes are a
region of the cerebral
cortex that is located
beneath the Sylvian
fissure on both the left
and right hemispheres of
the brain.
Lobes in this cortex are
more closely associated
with memory and in
particular
autobiographical
memory

The temporal lobes are


also concerned with
recognition memory.
This is the capacity to
identify an item as one that
was recently encountered.
Recognition memory is
widely viewed as consisting
of two components, a
familiarity component (i.e.
Do I know this person
waving at me?) and a
recollective component
(i.e. That is my friend Julia,
from evolutionary
psychology class).

Damage to the temporal


lobe can affect an
individual in a litany of
ways ranging from:
disturbance of auditory
sensation and perception,
disturbance of selective
attention of auditory and
visual input, disorders of
visual perception,
impaired organization
and categorization of
verbal material,
disturbance of language
comprehension, and
altered personality.

In regard to memory,
temporal lobe damage
can impair long-term
memory
Thus, general
semantic knowledge
or more personal
episodic memories of
ones childhood could
be affected.

The temporal lobe is involved in


auditory perception and is home to
the primary auditory cortex. It is also
important for the processing of
semantics in both speech and vision.
The temporal lobe contains the
hippocampus and plays a key role in
the formation of long-term memory.

The superior temporal gyrus includes an area (within


the Sylvian fissure) where auditory signals from the
cochlea (relayed via several subcortical nuclei) first
reach the cerebral cortex. This part of the cortex (
primary auditory cortex) is involved in hearing.
Adjacent areas in the superior, posterior and lateral
parts of the temporal lobes are involved in high-level
auditory processing. In humans this includes speech,
for which the left temporal lobe in particular seems to
be specialized. Wernicke's area, which spans the
region between temporal and parietal lobes, plays a
key role (in tandem with Broca's area, which is in the
frontal lobe). The functions of the left temporal lobe
are not limited to low-level perception but extend to
comprehension, naming, verbal memory and other
language functions.

The underside (ventral) part of the temporal cortices


appear to be involved in high-level visual processing of
complex stimuli such as faces (fusiform gyrus) and
scenes (parahippocampal gyrus). Anterior parts of this
ventral stream for visual processing are involved in
object perception and recognition.
The medial temporal lobes (near the Sagittal plane
that divides left and right cerebral hemispheres) are
thought to be involved in episodic/declarative memory.
Deep inside the medial temporal lobes lie the
hippocampi, which are essential for memory function particularly the transference from short to long term
memory and control of spatial memory and behavior.
Damage to this area typically results in
anterograde amnesia.

The occipital lobe is the visual processing center of


the mammalian brain containing most of the anatomical
region of the visual cortex.[1] The primary visual cortex
is Brodmann area 17, commonly called V1 (visual one).
Human V1 is located on the medial side of the occipital
lobe within the calcarine sulcus; the full extent of V1
often continues onto the posterior pole of the occipital
lobe. V1 is often also called striate cortex because it can
be identified by a large stripe of myelin, the
Stria of Gennari. Visually driven regions outside V1 are
called extrastriate cortex. There are many extrastriate
regions, and these are specialized for different visual
tasks, such as visuospatial processing, color
discrimination and motion perception. The name derives
from the overlying occipital bone, which is named from
the Latin oc- + caput, "back of the head".

Significant functional aspects of the occipital lobe is that it contains the


primary visual cortex and is the part of the brain where dreams come from.
Retinal sensors convey stimuli through the optic tracts to the
lateral geniculate bodies, where optic radiations continue to the visual
cortex. Each visual cortex receives raw sensory information from the
outside half of the retina on the same side of the head and from the inside
half of the retina on the other side of the head. The cuneus (Brodmann's
area 17) receives visual information from the contralateral superior retina
representing the inferior visual field. The lingula receives information from
the contralateral inferior retina representing the superior visual field. The
retinal inputs pass through a "way station" in the lateral geniculate nucleus
of the thalamus before projecting to the cortex. Cells on the posterior
aspect of the occipital lobes' gray matter are arranged as a spatial map of
the retinal field. Functional neuroimaging reveals similar patterns of
response in cortical tissue of the lobes when the retinal fields are exposed
to a strong pattern.
If one occipital lobe is damaged, the result can be homonomous vision loss
from similarly positioned "field cuts" in each eye. Occipital lesions can
cause visual hallucinations. Lesions in the parietal-temporal-occipital
association area are associated with color agnosia, movement agnosia,
and agraphia.

Parietal lobe
The parietal lobe is
located directly behind the
central sulcus, superior to
the occipital lobe and
posterior to the frontal
lobe, visually at the top of
the back of the head.
The make up of the
parietal lobe is defined by
four anatomical
boundaries in the brain,
providing a division of all
the four lobes.

The parietal lobe has


many functions and
duties in the brain and
its main functioning
can be divided down
into two main areas:
(1) sensation and
perception
(2) constructing a
spatial coordinate
system to represent
the world around us.

The parietal lobe helps us to


mediate attention when
necessary and provides spatial
awareness and navigational
skills. Also, it integrates all of our
sensory information (touch,
sight, pain etc.) to form a single
perception.
Parietal lobe gives the ability to
focus our attention on different
stimuli at the same time, PET
scans show high activity in the
parietal lobe when participates
being studied were asked to
focus their attention at two
separate areas of attention.
Parietal lobe also assists with
verbal short term memory and
damage to the supramarginal
gyrus cause short term memory
loss.

Damage to the parietal lobe


results in the syndrome
neglect which is when
patients treat part of their
body or objects in their visual
field as though it never
existed. Damage to the left
side of the parietal lobe can
result in what is called
Gerstmann syndrome
It includes right-left confusion,
difficulty with writing
(agraphia, and difficulty with
mathematics (acalculia. It can
also produce disorders of
language (aphasia, and the
inability to perceive objects

Damage to the right parietal


lobe can result in neglecting
part of the body or space
(contralateral neglect), which
can impair many self-care skills
such as dressing and washing.
Right side damage can also
cause difficulty in making things
(constructional apraxia, denial
of deficits (anosagnosia) and
drawing ability.
Neglect syndrome tends to be
more prevalent on the right side
of the parietal lobe, because the
right mediates attention to both
the left and right fields.
Damage in the somatic sensory
cortex results in loss of
perception of bodily sensations,
namely sense of touch.

The parietal lobe integrates sensory


information from different modalities,
particularly determining spatial sense and
navigation. For example, it comprises
somatosensory cortex and the dorsal stream
of the visual system. This enables regions of
the parietal cortex to map objects perceived
visually into body coordinate positions.
The name derives from the overlying
parietal bone, which is named from the Latin
pariet-, wall.

The parietal lobe plays important roles in integrating sensory information from
various parts of the body, knowledge of numbers and their relations, [1] and in the
manipulation of objects. Portions of the parietal lobe are involved with visuospatial
processing. Although multisensory in nature, the posterior parietal cortex is often
referred to by vision scientists as the dorsal stream of vision (as opposed to the
ventral stream in the temporal lobe). This dorsal stream has been called both the
'where' stream (as in spatial vision) [2] and the 'how' stream (as in vision for action).
[3]

Various studies in the 1990s found that different regions of the posterior parietal
cortex in Macaques represent different parts of space.
The lateral intraparietal (LIP) contains a map of neurons (retinotopically-coded
when the eyes are fixed[4]) representing the saliency of spatial locations, and
attention to these spatial locations. It can be used by the oculomotor system for
targeting eye movements, when appropriate. [5]
The ventral intraparietal (VIP) area receives input from a number of senses (visual,
somatosensory, auditory, and vestibular [6]). Neurons with tactile receptive fields
represented space in a head-centered reference frame. [6] The cells with visual
receptive fields also fire with head-centered reference frames [7] but possibly also
with eye-centered coordinates[6]
The medial intraparietal (MIP) area neurons encode the location of a reach target in
nose-centered coordinates.[8]
The anterior intraparietal (AIP) area contains neurons responsive to shape, size, and
orientation of objects to be grasped [9] as well as for manipulation of hands
themselves, both to viewed [9] and remembered stimuli.[10]

Occipital lobe

The occipital lobe is the smallest of


all four lobes in the human cerebral
cortex and located in the rearmost
part of the skull and considered to
be part of the forebrain
The occipital lobe sits directly above
the cerebellum and is situated
posterior to the Parieto-occipital
sulcus, or parieto-occipital sulcus.
This lobe is known as the centre of
the visual perception system, the
main function of the occipital lobe is
that of vision.

Retinal sensors send signals through


the optic tract to the Lateral
geniculate nucleus
Once the Lateral Geniculate Nucleus
receives the information it is sent
down the primary visual cortex
where it is organized and sent down
one of two possible path ways;
dorsal or ventral stream
The ventral stream is responsible for
object representation and
recognition and is also commonly
known as the "what" stream.
The dorsal stream is responsible for
guiding our actions and recognizing
where objects are in space,
commonly known as the "where" or
"how" stream.
Once in the information is organized
and send through the pathways it
continues to the other areas of the
brain responsible for visual
processing.

The most important function of the


Occipital lobe is vision.
Due to the positioning of this lobe at
the back of the head it is not
susceptible to much injury but any
significant damage to the brain can
cause a variety of damage to our
visual perception system.
Common problems in the occipital
lobe are field defects and scotomas,
movement and colour
discrimination, hallucinations,
illusions, inability to recognize
words and inability to recognize
movement.
A study was done in which patients
suffered from a tumour on the
occipital lobe and the results shows
that the most frequent consequence
was contralateral damage to the
visual field.

When damage occurs in the occipital


lobe it is most common to see the effects
on the opposite side of the brain. Since
the brain regions are so specialized in
their functioning damages done to
specific areas of the brain can cause
specific type of damage.
Damage to the left side of the brain can
lead to language discrepancies, i.e.
difficulty in properly identifying letters,
numbers and words, inability to
incorporate visual stimuli to comprehend
multiple ways an object can be found.

Right side damage causes non-verbal


problems, i.e. identifying geometric
shapes, perception of figures and faces

In almost all regions of the brain left side


damage leads to general language
problems where as right side damage
leads to general perception and problem
solving skills.

Damage to the cortex

Many studies of different


disease and disorders that
have symptoms of memory
loss have provided reinforcing
evidence to the study of the
anatomy of the brain and
which parts are more utilized
in memory.

Frontotemporal lobar
degeneration (FTLD) is a
common form of dementia due to
the degeneration of the frontal
and temporal lobes. Studies have
found significant decreases in the
essential needs for proper
functioning in these lobes.
The autobiographical domain in
memory is largely affected by
this disease. In one study, FTLD
patients were interviewed and
asked to describe a significant
event from five different periods
of their lives.
Using the interview and different
methods of imaging, the
experimenters hoped to find links
between patterns of brain
volume loss and performance in
the interview

Parkinson's disease

involves both damage to the


basal ganglia and certain
memory dysfunctions,
suggesting that the basal
ganglia are involved in specific
types of memory.
Those who have this disease
have problems with both their
working memory and spatial
memory.
Most people can instantly and
easily use visual-spatial
memory to remember locations
and pictures, but a person with
Parkinson's disease would find
this difficult.
He or she would also have
trouble encoding this visual and
spatial information into longterm memory.

This suggests that the basal


ganglia work in both encoding
and recalling spatial
information.
People with Parkinson's
disease display working
memory impairment during
sequence tasks and tasks
involving events in time.
They also have difficulty in
knowing how to use their
memory, such as when to
change strategies or maintain
a train of thought.

Recall: neuroanatomy

The anterior cingulate cortex,


globus pallidus, thalamus, and
cerebellum show higher
activation during recall than
during recognition which
suggests that these
components of the cerebellofrontal pathway play a role in
recall processes that they do
not in recognition. Although
recall and recognition are
considered separate
processes, it should be noted
that they are both most likely
constitute components of
distributed networks of brain
regions.[

According to neuroimaging data,


PET studies on recall and
recognition have consistently
found increases in regional
cerebral blood flow (RCBF) in the
following six brain regions:
(1) the prefrontal cortex
particularly on the right
hemisphere; (2) the hippocampal
and parahippocampal regions of
the medial temporal lobe; (3) the
anterior cingulate cortex; (4) the
posterior midline area that
includes posterior cingulate,
retrosplenial (see retrosplenial
region), precuneus, and cuneus
regions; (5) the inferior parietal
cortex, especially on the right
hemisphere; and (6) the
cerebellum, particularly on the
left.[

Factors that affect recall

Attention
The effect of attention on memory
recall has surprising results. It
seems that the only time attention
largely affects memory is during the
encoding phase . During this phase,
performing a parallel task can
severely impair retrieval success.
It is believed that this phase
requires much attention to properly
encode the information at hand, and
thus a distractor task does not allow
proper input and reduces the
amount of information learned.

Motivation
Motivation is a factor that
encourages a person to perform and
succeed at the task at hand. It can
be in the form of presented
incentive, or personal fear of failure.
Any form of motivation thus
generally leads a person to better
recall. In an experiment done by
Roebers, Moga and Schneider
(2001), participants were placed in
either forced report, free report or
free report plus incentive groups. In
each group, they found that the
amount of correct information
recalled did not differ, yet in the
group where participants were given
an incentive they had higher
accuracy results.
This means that presenting
participants with an encouragement
to provide correct information
motivates them to be more precise.

Interference
In the absence of interference, there are two
factors at play when recalling a list of items:
the recency and the primacy effects. The
recency effect occurs when the short-term
memory is used to remember the most recent
items, and the primacy effect occurs when
the long-term memory has encoded the
earlier items. The recency effect can be
eliminated if there is a period of interference
between the input and the output of
information extending longer than the holding
time of short-term memory (1530 seconds).
This occurs when a person is given
subsequent information to recall preceding
the recall of the initial information.[48] The
primacy effect, however, is not affected by
the interference of recall. The elimination of
the last few items from memory is due to the
displacement of these items from short term
memory, by the distracting task. As they have
not been recited and rehearsed, they are not
moved into long-term memory and are thus
lost. A task as simple as counting backwards
can change memory recall; however an
empty delay interval has no effect.[49] This is
because the person can continue to rehearse
the items in their working memory to be
remembered without interference

Context
Context-dependency effects on recall are
typically interpreted as evidence that the
characteristics of the environment are
encoded as part of the memory trace and
can be used to enhance retrieval of the
other information in the trace.[52] In other
words, you can recall more when the
environments are similar in both the
learning and recall phases. Context cues
appear to be important in the retrieval of
newly learned meaningful information. In
a classic study by Godden and Baddelley
(1975), they demonstrated that deep-sea
divers recalled their training more
effectively when trained underwater,
rather than being trained on land.[53] An
academic application would be that
students may perform better on exams by
studying in silence, because exams are
usually done in silence

State-dependent memory
State-dependent retrieval is demonstrated
when material learned under the influence of
a drug is best recalled in that same drug
state. A study by Carter and Cassady (1998)
showed this effect with antihistamine.[55] In
other words, if you study while on hay fever
tablets, then you will recall more of what you
studied if you test yourself while on
antihistamines in comparison to testing
yourself while not on antihistamines after
having studied on antihistamines.
A study by Block and Ghoneim (2000) found
that, relative to a matched group of healthy,
non-drug-using controls, heavy marijuana use
is associated with small but significant
impairments in memory retrieval.[56]Cannabis
induces loss of internal control and cognitive
impairment, especially impairment of
attention and memory, for the duration of the
intoxication period.[57]
Stimulants, such as cocaine, amphetamines
or caffeine are known to improve recall in
humans.[58] However, the effect of prolonged
use of stimulants on cognitive functioning is
very different from the impact on one-time
users. Some researchers have found
stimulant use to lower recall rates in humans
after prolonged usage

Gender
Consistently, females perform better than
males on episodic memory tasks including
delayed recall and recognition. However,
males and females do not differ on working,
immediate and semantic memory tasks. In
general, neuro-psychological observations
suggest that anterior lesions cause greater
deficits in females than in male. It has been
proposed that the gender differences in
memory performance reflect underlying
differences in the strategies used to process
information, rather than anatomical
differences. However, gender differences in
cerebral asymmetry received support from
morphometric studies showing a greater
leftward asymmetry in males than in
females, meaning that men and women use
each side of their brain to a different
extent.[59] There is also evidence for a
negative recall bias found in women, which
means females in general are more likely
than males to recall their mistakes.[60] In a
eyewitness study done by Dan Yarmey
(1991) from the University of Guelph, he
found that women were significantly more
accurate than men in accuracy of recall for
weight of suspects

Phenomena
The phenomenological account
of recall is referred to as
metacognition, or "knowing
about knowing". This includes
many states of conscious
awareness known as feeling-ofknowing states, such as the tipof-the-tongue state. It has been
suggested that metacognition
serves a self-regulatory purpose
whereby the brain can observe
errors in processing and actively
devote resources to resolving
the problem. It is considered an
important aspect of cognition
that can aid in the development
of successful learning strategies
that can also be generalized to
other situations.[62]

The cerebellum
(Latin for little brain) is a
region of the brain that plays
an important role in
motor control. It is also
involved in some
cognitive functions such as
attention and language, and
probably in some emotional
functions such as regulating
fear and pleasure responses.[
Its movement-related
functions are the most clearly
understood, however.
The cerebellum does not
initiate movement, but it
contributes to coordination,
precision, and accurate timing.

It receives input from


sensory systems and from
other parts of the brain
and spinal cord, and
integrates these inputs to
fine tune motor activity.[
Because of this fine-tuning
function, damage to the
cerebellum does not cause
paralysis, but instead
produces disorders in fine
movement, equilibrium,
posture, and
motor learning

cerebellu
m

Cerebellum

The cerebellum
The cerebellum ("little brain") is a
structure located at the rear of the
brain, near the spinal cord
It looks like a miniature version of
the cerebral cortex, in that it has a
wavy, or convoluted surface.

Unlike the hippocampus which is


involved in the encoding of complex
memories, the cerebellum plays a
role in the learning of procedural
memory, and motor learning, such
as skills requiring co-ordination and
fine motor control.
An example of a skill requiring
procedural memory would be
playing a musical instrument, or
driving a car or riding a bike.
Individuals with transient global
amnesia that have difficulty forming
new memories and/or remembering
old events may sometimes retain
the ability to perform complex
musical pieces, suggesting that
procedural memory is completely
dissociated from conscious memory,
also known as explicit memory

This separation
makes sense if the
cerebellum, which is
far removed from
the hippocampus, is
responsible for
procedural learning.

The cerebellum is more


generally involved in
motor learning, and
damage to it can result
in problems with
movement, specifically
it is considered to coordinate timing and
accuracy of
movements, and to
make long-term
changes (learning) to
improve these skills

Based on surface appearance,


three lobes can be
distinguished in the
cerebellum, called the
flocculonodular lobe,
anterior lobe (above the
primary fissure), and
posterior lobe (below the
primary fissure). ")

These lobes divide the


cerebellum from rostral to
caudal (in humans, top to
bottom).
In terms of function, however,
there is a more important
distinction along the medial-tolateral dimension.
Leaving out the flocculonodular
part, which has distinct
connections and functions, the
cerebellum can be parsed
functionally into a medial sector
called the spinocerebellum and
a larger lateral sector called the
cerebrocerebellum.
A narrow strip of protruding
tissue along the midline is called
the vermis (Latin for "worm

The smallest region, the


flocculonodular lobe, is often
called the
vestibulocerebellum. It is the
oldest part in evolutionary
terms (archicerebellum) and
participates mainly in balance
and spatial orientation; its
primary connections are with
the vestibular nuclei, although
it also receives visual and
other sensory input. Damage
to it causes disturbances of
balance and gait.[

The medial zone of the anterior


and posterior lobes constitutes
the spinocerebellum, also
known as paleocerebellum.
This sector of the cerebellum
functions mainly to fine-tune
body and limb movements.
It receives proprioception input
from the dorsal columns of the
spinal cord (including the
spinocerebellar tract) and from
the trigeminal nerve, as well as
from visual and auditory
systems.
It sends fibres to deep
cerebellar nuclei that, in turn,
project to both the cerebral
cortex and the brain stem, thus
providing modulation of
descending motor systems.

The lateral zone, which in


humans is by far the largest
part, constitutes the
cerebrocerebellum, also
known as neocerebellum.
It receives input exclusively
from the cerebral cortex
(especially the parietal lobe)
via the pontine nuclei
(forming cortico-pontocerebellar pathways), and
sends output mainly to the
ventrolateral thalamus (in
turn connected to motor
areas of the premotor
cortex and primary motor
area of the cerebral cortex)
and to the red nucleus.

There is disagreement
about the best way to
describe the functions of
the lateral cerebellum: It is
thought to be involved in
planning movement that is
about to occur, in
evaluating sensory
information for action, and
in a number of purely
cognitive functions as well.

Prior to the 1990s, the function


of the cerebellum was almost
universally believed to be
purely motor-related, but
newer findings have brought
that view strongly into
question. Functional imaging
studies have shown cerebellar
activation in relation to
language, attention, and
mental imagery; correlation
studies have shown
interactions between the
cerebellum and non-motoric
areas of the cerebral cortex;
and a variety of non-motor
symptoms have been
recognized in people with
damage that appears to be
confined to the cerebellum.

The cerebellum, Doya


proposes, is best
understood as a device
for supervised
learning, in contrast to
the basal ganglia,
which perform
reinforcement learning,
and the cerebral
cortex, which performs
unsupervised learning.

There is
considerable
evidence that the
cerebellum plays
an essential role in
some types of
motor learning.

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