Biological Foundations of Behaviour – I

Unit 3 - Biopsychology of Cognitive

Functions
Learning
� Learning creates new information
� the process by which experiences change our nervous system and our
behavior
� Long-term changes leading to form memories.
� Memories form over new learning or relearning.
� Learning new and creating a memory - physically changes the structure of
the nervous system, altering neural circuits that participate in perceiving,
performing, thinking, planning, and behaving.
Process of Learning - The information-
processing model of memory
� New learning - produces changes in the nervous system - encoding the
new information to be learnt.
� The encoding process + consolidation + storage + Retrieval
� consolidation - strengthens changes associated with the initial information
being learnt, helps make permanent changes to the nervous system
� Storage - memory is stored via these persistent changes in the nervous
system.
� Retrieval is the process of accessing and using the information stored in
the neural changes that make up a memory to engage in a behavior.
 Types of Learning
� S-R Learning: Classical Conditioning & Operant
Conditioning
� Motor learning

� Perceptual learning

� Relational learning
Types of Memory
 Lashley’s Search for the engram
� Karl Lashley
� “engram”—permanent change in the brain accounting for the
existence of memory; a memory trace - the physical representation of
what has been learned - A connection between two brain areas would
be a possible example of an engram.
� if learning depends on new or strengthened connections between two
brain areas, a knife cut somewhere in the brain should interrupt that
connection and abolish the learned response.
� He trained rats on mazes and a brightness discrimination task and
then made deep cuts in varying locations in their cerebral cortices - no
knife cut significantly impaired the rats’ performances - the types of
learning that he studied did not depend on connections across the
cortex.
� Lashley also tested whether any portion of the cerebral cortex is
more important than others for learning - He trained rats on mazes
before or after he removed large portions of the cortex - The lesions
impaired performance, but the deficit depended more on the amount
of brain damage than on its location - Learning and memory
apparently did not rely on a single cortical area.
� therefore proposed two principles about the nervous system:
� equipotentiality—all partsof the cortex contribute equally to
complex behaviors such as learning, and any part of the cortex can
substitute for any other.
� mass action—the cortex works as a whole, and more cortex is
better
� Richard F. Thompson and his colleagues used a simpler task
seeking the location of engram.
� sought the engram of memory not in the cerebral cortex but in the
cerebellum.
� studied classical conditioning of eyelid responses in rabbits.
They presented first a tone (CS) and then a puff of air (UCS) to
the cornea of the rabbit’s eye. At first, a rabbit blinked at the air
puff but not at the tone. After repeated pairings, classical
conditioning occurred and the rabbit blinked at the tone also.
� Investigators recorded the activity in various brain cells to
determine which ones changed their responses during learning.
� Thompson - determined the location of learning.
� sequence of brain areas from the sensory receptors to the motor neurons
controlling the muscles – a damage any one of those areas, learning will
be impaired, but we cannot be sure that learning occurred in the damaged
area.
� For example, if the learning occurs in area D, damage in A, B, or C will
prevent learning by blocking the input to D. Damage in E or F will
prevent learning by blocking the output from D.
� Thompson and colleagues reasoned as follows: Suppose the learning
occurs in D. If so, then D has to be active at the time of the learning, and
so do all the areas leading up to D (A, B, and C). However, learning
should not require areas E and beyond. If area E were temporarily
blocked, nothing would relay information to the muscles, so we would see
no response, but learning could occur nevertheless, and we could see
evidence of it later
� Thompson’s research identified one nucleus of the cerebellum - the lateral
interpositus nucleus (LIP) - essential for learning.
� At the start of training, those cells showed little response to the tone, but as
learning proceeded, their responses increased
� When temporarily suppressed that nucleus in an untrained rabbit, either by
cooling the nucleus or by injecting a drug into it, and then presented the
CSs and UCSs, the rabbit showed no responses during the training. Then
they waited for the LIP to recover and continued training. At that point, the
rabbit began to learn, but it learned at the same speed as animals that had
received no previous training. Evidently, while the LIP was suppressed, the
training had no effect
� The next question was - does learning actually occur in the LIP, or does this
area just relay the information to a later area where learning occurs?
� To understand - investigators suppressed activity in the red nucleus, a
midbrain motor area that receives input from the cerebellum.
� When the red nucleus was suppressed, the rabbits again showed no
responses during training. However, as soon as the red nucleus had
recovered from the cooling or drugs, the rabbits showed strong learned
responses to the tone
� In other words, suppressing the red nucleus temporarily prevented the
response but did not prevent learning.
� Evidently, learning did not require activity in the red nucleus or any area
after it, although later research found that the red nucleus does contribute to
learning under certain circumstances
� Thompson and his colleagues concluded – “the learning occurred in the
LIP”.
Lashley and � The mechanisms for this type of conditioning are probably
Thompson in similar in humans.
Human � According to PET scans on young adults - developing a
learning conditioned eyeblink causes increases in the cerebellum, red
nucleus, and several other areas
� People who have damage in the cerebellum have weaker
conditioned eyeblinks, and the blinks are less accurately timed
relative to the onset of the air puff
� The cerebellum is critical for many other instances of classical
conditioning also, but only if the delay between the onset of the
CS and the onset of the UCS is short.
� the cerebellum is specialized for timing brief intervals, on the
order of a couple of seconds or less.
� Many instances of learning take place in other brain areas. For
example, learning to avoid a taste because of subsequent illness
depends on the amygdala
 Classical Conditioning
ROLE OF AMYGDALA
• The amygdala is important in classically conditioned emotional responses
• Eg:- classically conditioned emotional response is established by pairing a neutral
stimulus (such as a tone) with an aversive stimulus (such as a brief foot shock) - after
these stimuli are paired, the tone becomes a CS; when it is presented by itself, it elicits the
same type of responses as the unconditioned stimulus does.
• After being processed by the auditory cortex, information about the CS (the tone) reaches
the lateral nucleus of the amygdala. This nucleus also receives information about the US
(the foot shock) from the somatosensory system. Thus, these two sources of information
converge in the lateral nucleus, which means that synaptic changes responsible for
learning could take place in this location
Auditory cortex – lateral nucleus of Amygdala + somatosensory system = converge in
lateral nucleus – changes in synaptic
� The lateral nucleus of the amygdala contains neurons whose axons project
to the central nucleus.
� Terminal buttons from neurons that transmit auditory and somatosensory
information to the lateral nucleus form synapses with dendritic spines on
these neurons - When a rodent encounters a painful stimulus, somatosensory
input activates strong synapses in the lateral nucleus.
� As a result, the neurons in this nucleus begin firing, which activates neurons
in the central nucleus, evoking an unlearned (unconditioned) emotional
response. If a tone is paired with the painful stimulus, the weak synapses in
the lateral amygdala are strengthened. The synaptic changes responsible for
this type of learning take place within this circuit.
Study on
Aplysia
ROLE OF GLUTAMATE
� the lateral amygdala responsible for acquisition of a conditioned
emotional response involve a series of synaptic changes called long-term
potentiation (LTP). LTP involves NMDA (N-methyl-D-aspartate) receptors
and AMPA (amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid) glutamate
receptors.
� LTP is accomplished through the activation of NMDA receptors and the
insertion of additional AMPA receptors into the postsynaptic membrane.
These synaptic changes in the glutamate system serve to increase the
Excitatory Post Synaptic Potential (EPSP) to the postsynaptic cell.
� LTP among glutamate synapses in the lateral amygdala plays a critical
role in the establishment of conditioned emotional responses.
� The AMPA opens🡪 let the ions enter
inside-depolarizes the neurons🡪 results in
the eviction of Mg2+ from the NMDA
receptors that blocks it(due to electrostatic
force)🡪 NMDA receptors open🡪 Let the
Ca2+ inside🡪 triggers enzyme synthesis
🡪 triggers the AMPA receptors from the
other regions of the cell membrane to move
and insert on the postsynaptic dendritic
spines🡪 More glutamates can now bind
with the number of receptors increases🡪
results in eventual increased EPSP
(Excitatory postsynaptic potentiation)🡪
results in stronger synaptic
connection(neuroplasticity)
Operant Conditioning- ROLE OF BASAL GANGLIA

� The circuits responsible for OC - begin in various regions of the sensory

association cortex, where perception takes place, and end in the motor
association cortex of the frontal lobe, which controls movements.
� There are two major pathways between the sensory association cortex and
the motor association cortex
� direct transcortical connections - connections from one area of the
cerebral cortex to another
� Basal Ganglia pathway - connections via the basal ganglia and
thalamus.
Transcortical Pathways –
� along with the hippocampal formation, the transcortical connections are
involved in the acquisition of declarative, episodic memories—complex
perceptual memories of sequences of events that we experience or that are
described to us.
� also involved in the acquisition of complex behaviors that involve deliberation
or instruction – eg. Instructor’s instructions while driving a car.
� At first – rules and processes make it difficult to perform an activity, but with
practice / repeated exposure – it is easy to perform the activity.
Basal Ganglia Pathways
� Evidence suggests that as learned behaviors become automatic and routine, -
they are “transferred” to the basal ganglia.
� The process - As we deliberately perform a complex behavior, the basal
ganglia receive information about the stimuli that are present and the
responses we are making. At first the basal ganglia are passive “observers” of
the situation, but as the behaviors are repeated again and again, the basal
ganglia begin to learn what to do. Eventually, they take over most of the
details of the process, leaving the transcortical circuits free to do something
else. We need no longer think about what we are doing. Returning to the
example of driving a car with a manual transmission, as the driver practices,
processing is transferred to the basal ganglia, and the driver no longer needs to
deliberately think through each step. Instead, the driver fluidly and
automatically starts the engine, shifts gears, and drives the car
 Reinforcement-Role of Dopamine
� Experiment by James Olds and Peter
Milner (1954) on rats pioneered the
investigation in this area
� They discovered that it is the
dopaminergic neurons in the ventral
tegmental area (VTA-part of midbrain)
plays crucial role in reinforcement.
� VTA is connected to amygdala,
hippocampus and Nucleus accumbens
(NAC)
� Stimulation to medial forebrain bundle
(MFB) or VAT causes the release of
dopamine in the NAC.
� Operant conditioning involves three elements-discriminative stimulus, response
and reinforcement.
� Only when a behaviour/response is followed by a reinforcement will triggers the
secretion od dopamine
� Experiment by Knetch et al., (2004)-learning artificial vocabulary and subsequent
administration of L-DOPA (precursor of dopamine) resulted in greater memory of
the words
 Detecting reinforcement-Expected vs Unexpected
� Prefrontal cortex provides inputs to
VTA, results in glutamate secretion.
� This activity triggers the
dopaminergic neurons in the VTA to
secrete more dopamine in the NAC.
� As the prefrontal cortex involved in
the planning, evaluation and
judgement of the behaviour, it may
turn on the reinforcement
mechanism.
Motor Learning-Role of cortical and subcortical structures
� Motor learning typically involves learning a novel sequence of motor
behaviors over repeated trials. The cerebellum, thalamus, basal ganglia,
and motor cortex are involved in motor learning across many different
tasks
� Motor behaviours maybe either reflexive or a series of actions that
needs to be learned and later becomes automatic.
� Several cortical and subcortical regions are simultaneously involved in
sequential motor activities.
� Cortical structures involved in the control of movement include the
primary motor cortex, supplementary motor area, and the
premotor cortex.
 Cortical structures

� Cortical structures involved in

the control of movement
include the primary motor
cortex, supplementary motor
area, and the premotor cortex.
� Study by Graziano and Aflalo (2007) on
monkeys (p.no: 256, Carlson)
� Brief stimulation to of the particular
primary motor cortex resulted in the
brief movements of various part of the
body, whereas the prolonged stimulation
caused much more complex movements.
� Supplementary Motor Area and
Premotor Cortex receive sensory
information from the parietal and
temporal lobes, and both send efferent
axons to the primary motor cortex.
 Subcortical structures

� motor behavior also involves several subcortical

structures, including the reticular formation,
cerebellum, and the basal ganglia.
� These structures contain sets of nuclei in the
mid- and hindbrain that are involved in control
of voluntary and involuntary motor behavior,
posture, locomotion, and limb movements.
� They receive information from regions of the
motor cortex and help convey it to the spinal
cord and nerves.
Reticular formation
� consists of a large number of nuclei located in the core of the medulla, pons, and midbrain.
� It plays a role in the control of posture and locomotion.
Cerebellum
� Flocculonodular lobe, located at the caudal end of the cerebellum, receives input from the vestibular
system and projects axons to the vestibular nucleus. It is involved in the postural reflexes.
� The vermis--🡪 located on the midline, receives auditory and visual information from the tectum
and cutaneous and kinesthetic information from the spinal cord. It sends its outputs to the fastigial
nucleus (one of the set of deep cerebellar nuclei). Neurons in the fastigial nucleus send axons to the
vestibular nucleus and to motor nuclei in the reticular formation (RF). Thus, these neurons influence
behavior through the vestibulospinal and reticulospinal tracts, two of the three ventromedial pathways.
Vermis🡪 Fastigial nucleus🡪 Vestibular nucleus (medulla) and motor nuclei (RF)
� Rest of the inputs to the cerebellum comes from the cerebral cortex
Cerebral cortex🡪 Pontine tegmental reticular nucleus🡪 LIP of cerebellum🡪 Red nucleus
The intermediate zone of the cerebellum influences the control of the rubrospinal system over movements
of the arms and legs.
The lateral zone of the cerebellum is involved in the control of independent limb movements, especially
rapid, skilled movements.
� Although the frontal cortex can plan and initiate movements, it does not contain the neural
circuitry needed to calculate the complex, closely timed sequences of muscular contractions that are
needed for rapid, skilled movements.
� That task falls to the lateral zone of the cerebellum. Both the frontal association cortex and the
primary motor cortex send information about intended movements to the lateral zone of the
cerebellum via the pontine nucleus.
� The lateral zone also receives information from the somatosensory system, which informs it about
the current position and rate of movement of the limbs—information that is necessary for
computing the details of a movement. When the cerebellum receives information that the motor
cortex has begun to initiate a movement, it computes the contribution that various muscles will
have to make to perform that movement.
� The results of this computation are sent to the dentate nucleus, another of the deep cerebellar
nuclei. Neurons in the dentate nucleus pass the information on to the ventrolateral thalamus, which
projects to the primary motor cortex.
� The projection from the ventrolateral thalamus to the primary motor cortex enables the cerebellum
to modify the ongoing movement that was initiated by the frontal cortex.
� The lateral zone of the cerebellum also sends efferents to the red nucleus (again, via the dentate
nucleus); thus, it helps to control independent limb movements through this system as well.
 In sum,
Red nucleus

Frontal
association cortex
Pontine lateral zone Dentate
ventrolater Primary
nucleus of of nucleus of
tegmentum cerebellum cerebellum al thalamus motor cortex
primary motor
cotex

Somatosensory
system
Basal ganglia
• Direct pathway- Excitatory
• Indirect pathway- Inhibitory
• Hyper direct pathway-Quick inhibitory
Role of Mirror Neurons in imitating and comprehending movements

Refer to Carlson-P. no:269

 Perceptual Learning
� Perceptual learning involves learning to recognize things. It can involve learning to
recognize entirely new stimuli, or it can involve learning to recognize changes or
variations in familiar stimuli.
� Eg: Our visual memory of people changes when there is a change in their physical
appearance
� This type of learning is actually involved in other types of learning as well as it helps
in identifying the stimuli.
� The location or the context of the stimuli also matters in their recognition
� Learning and remembering various stimuli involve several cortical mechanisms
� Memory also involves complex series of events or episodes that involves numerous
contextually relevant stimuli.
� This section covers the role of cortex in learning to recognize stimuli,
remembering them later and retention of perceptual information in short-term
memory.
 Role of cortex in perceptual learning
� In mammals with large and complex brains, objects are recognized visually by circuits of
neurons in the extra-striate cortex.
� Visual learning can take place very rapidly, and the number of items that can be
remembered is enormous.
� Other primates are capable of remembering items that they have seen for just a few
seconds, and the experience changes the responses of neurons in their extrastriate cortex
(Rolls, 1995).
� Extrastriate cortex is the visual association cortex, whereas striate cortex is the primary
visual cortex (V1).
� Extra striate cortex involves different regions (V2 to V5) each analyse different aspects of
visual stimuli including the orientation, movement, color, retinal disparity etc. It is a
hierarchical pathway in which information from V1 goes to the successive levels of
extrastriate cortex, V2….V5.
� striate cortex receives information from the lateral
geniculate nucleus of the thalamus.
� After the first level of analysis in the striate cortex, the
information is sent to the extrastriate cortex.
� After analyzing particular attributes of the visual scene,
such as form, color, and movement, subregions of the
extrastriate cortex send the results of their analysis to
the next level of the visual association cortex
� This level is divided into two “streams.” The ventral
stream and dorsal stream.
� Ventral stream is involved with object recognition
(what), continues ventrally into the inferior temporal
cortex.
� The dorsal stream is involved with perception of the
location of objects (where), continues dorsally into the
posterior parietal cortex.
� lesions that damage the inferior temporal cortex—the end of the ventral
stream— disrupt the ability to discriminate among visual stimuli. These
lesions impair the ability to perceive (and thus to learn to recognize)
particular kinds of visual information. People with damage to the inferior
temporal cortex may have excellent vision but be unable to recognize
familiar, everyday objects such as scissors, cell phones, or light bulbs—
and faces of friends and relatives.
Role of cortex in remembering perceptual stimuli

� Perceptual learning involves changes in synaptic connections in the extrastriate cortex that
establish new neural circuits.
� At a later time, when the same stimulus is seen again and the same pattern of activity is
transmitted to the cortex, these circuits become active again.
� This activity constitutes the recognition of the stimulus—the readout, or replay, of the visual
memory.
� Yang and Maunsell (2004) trained monkeys to detect small differences in visual stimuli whose
images were projected onto a specific region of the retina. After the training was complete, the
monkeys were able to detect differences much smaller than those they could detect when the
training first started. However, they were unable to detect these differences when the patterns
were projected onto other regions of the retina. Recordings of single neurons in the extrastriate
cortex showed that the response properties of neurons that received information from the
“trained” region of the retina—but not from other regions—had become sensitive to small
differences in the stimuli. Neural circuits in that region alone had been modified by the training.
(https://www.jneurosci.org/content/24/7/1617.short)
� When stimulated the extrastriate and auditory association cortex as patients were
undergoing seizure surgery, the patients reported memories of images or sounds—for
example, images of a familiar street or the sound of the patient’s mother’s voice (seizure
surgery is performed under a local anesthetic so that the surgeons can test the effects of
brain stimulation on the patients’ cognitive functions).
(Penfield & Perot, 1963)
� Damage to regions of the brain involved in visual perception not only impairs the ability
to recognize visual stimuli but also disrupts people’s memory of the visual properties of
familiar stimuli.
� For example, Vandenbulcke et al., (2006) found that Patient J. A., who had sustained
damage to the right fusiform gyrus, performed poorly on tasks that required her to draw
or describe visual features of various animals, fruits, vegetables, tools, vehicles, or
pieces of furniture. Her other cognitive abilities, including the ability to describe
nonvisual attributes of objects, were intact. In addition, an fMRI study found that when
healthy individuals were asked to perform the visual tasks that she performed poorly,
activation was seen in the region of their brains that corresponded to J. A.’s lesion (in the
right fusiform gyrus).
� Kourtzi and Kanwisher (2000) found that specific kinds of visual information can
activate very specific regions of extrastriate cortex. A region of the extrastriate cortex,
MT/MST, plays an essential role in perception of movement.
� The investigators presented participants with photographs that implied motion—for
example, an athlete getting ready to throw a ball. They found that photographs like
these, but not photographs of people remaining still, activated area MT/MST. Even
though the photographs did not move, the participants’ memories presumably
contained information about movements they had previously seen.
� A functional-imaging study by Goldberg et al., (2006) asked people questions that
involved visual, auditory, tactile, and gustatory information. The researchers found that
answering the questions activated the regions of association cortex involved in
perception of the relevant sensory information. For example, questions about flavour
activated the gustatory cortex, questions about tactile information activated the
somatosensory cortex, and questions about visual and auditory information activated
the visual and auditory association cortex.
Retaining Perceptual Information in Short-Term Memory
� Most often we immediately after sensing a stimulus, we respond. But sometimes the situation
demands that we make the appropriate response after a delay, even after the stimulus is no longer
visible.
� For example: Crossing a road, parking a car
� Tasks like these demands comparing a perception with short term memory of something else we
had just perceived.
� Learning to recognize a stimulus involves synaptic changes in the appropriate regions of the
sensory association cortex that establish new circuits of neurons.
� Recognition of a stimulus occurs when sensory input activates these established sets of neural
circuits.
� Short-term memory of a stimulus involves activity of these circuits— or other circuits that are
activated by them—that continues even after the stimulus disappears.
� E.g.: learning to recognize a friend’s face produces changes in synaptic strengths in neural circuits
in the fusiform face region of our visual association cortex, recognizing that she is present involves
activation of the circuits that are established by these changes, and remembering that she is still in
the room even when we look elsewhere involves continued activity of these circuits (or other
circuits connected to them).
ROLE OF EXTRASTRIATE CORTEX

� Functional-imaging studies have shown that retention of specific types of short-term

visual memories involves activity of specific regions of the extrastriate cortex that we
encountered in Chapter 6. One region of the ventral stream, the fusiform face area, is
involved in recognition of faces, and another region, the parahippocampal place area (part
of the ventral stream), is involved in recognition of places. A functional-imaging study by
Ranganath et al., (2004) found evidence that short-term memory for particular faces and
places was associated with neural activity in two different regions of the ventral stream of
the visual association cortex. The investigators trained people on a delayed matching-to-
sample task that required them to remember particular faces or places for a short period of
time. In a delayed matching-to-sample task, a participant is shown a stimulus (the
sample), and then, after a delay during which the stimulus disappears, the participant must
indicate which of several alternatives matches the sample. Ranganath and his colleagues
found that short-term memories of faces activated the fusiform face area and that short-
term memories of places activated the parahippocampal place area.
� Transcranial magnetic stimulation (TMS) of the extrastriate cortex interferes with visual
perception. TMS can be used to induce a weak electrical current in the brain that disrupts
neural activity and thus interferes with the normal functions of the stimulated region.
Oliveri et al., (2001) trained people on a delayed matchingto-sample task that required
them to remember either abstract figures or the locations of a white square on a video
screen. On some trials the investigators applied TMS to regions of either the ventral stream
or the dorsal stream during the delay interval, after the sample stimuli had been turned off.
They found that, as expected, stimulating the ventral stream interfered with short-term
memory for visual patterns and stimulating the dorsal
ROLE OF PREFRONTAL CORTEX

� Although the neural circuits responsible for learning to recognize particular stimuli appear
to reside in the sensory association cortex, perceptual short-term memories involve other
brain regions as well—especially the prefrontal cortex. Miyashita (2004) suggests that the
role of the prefrontal cortex in short-term memory is to “manipulate and organize to-be-
remembered information, devise strategies for retrieval, and also monitor the outcome” of
these processes (p. 435).
� Baier et al., (2010) note that successfully remembering recently presented information in
short-term memory requires two processes: filtering out irrelevant information and
maintaining relevant information. The investigators presented stroke patients with short-
term memory tasks that either required them to ignore extraneous, irrelevant information
or tested their ability to hold several pieces of information in mind. They found that
patients with damage to the left basal ganglia had difficulty filtering out irrelevant
information, and patients with damage to the right prefrontal cortex had difficulty retaining
more
Relational Learning
� Most of the real life learnings more complex and the memories of
objects and events are inter-related.
� Remembering an object of event may lead to retrieval of a series of
connected objects and events.
� Neural circuits in the visual cortex, especially extrastriata cortex
plays a crucial role in the recognition of a stimuli.
� These circuits are connected to the circuits in the other parts of the
brain, they are connected to the other and so on.
 Role of the Hippocampus in relational learning
� Hippocampal formation consists of dentate gyrus, regions called the CA fields of the hippocampus,
and the subiculum.
� Inputs from
Amygdala Field CA1
Entorhinal
Regions of limbic cortex Entorhinal cortex Perirhinal Associati
all association regions of cortex Subiculum parahippocampal on cortex

Perirhinal & parahippocampal

Cortex
Inputs to the entorhinal cortex can be either directly from amygdala, regions of limbic cortex and all
regions of association cortex or via Perirhinal & parahippocampal Cortex (two adjacent regions of limbic cortex)
A closer look into all these structures
� Hvbn
� Gfh
� Nbvcv
Hippocampal formation also receives inputs form the subcortical regions via fornix.
Fornix connects hippocampal formation with the mammillary bodies
The input travels from
Mammillary
Thalamus
bodies

Ventral tegmental area

Locus coeruleus Hippocampal
Fornix
Raphe nuclei formation
Medial septum

Degeneration of mammillary bodies is implicated in the Korsakoff’s syndrome and

maybe in anterograde amnesia.
� Vbv
 Consolidation of memories-Role of hippocampus
� The hippocampal formation plays a role in the process through which declarative memories are formed.
� The hippocampus receives information from cortical and subcortical regions.

Hippocampus’ efferent connections with Sensory and

Sensory and motor motor association cortex, basal ganglia, amygdala
association cortex, Hippocampus modifies the memories that are being consolidated
basal ganglia, there, linking them together in ways that will permit
amygdala us to remember the relationships among the elements
of the memories

� Without the hippocampal formation, we would be left with individual, isolated memories without
the linkage that makes it possible to remember—and think about—episodes and contexts.
� Acquisition of both major categories of relational, declarative memories—episodic and
semantic—appears to require the participation of the hippocampus. Manns et al., (2003) found
that five patients with damage limited to the hippocampal formation showed an anterograde
amnesia for semantic as well as episodic information.
� Perceptual memories appear to be located in the sensory association cortex, the
regions where the perceptions take place. Presumably, episodic memories, which
consist of an integrated sequence of perceptual memories, are also located there.
� However, the semantic memories are not simply perceptual memories. A
degenerative neurological disorder known as semantic dementia suggests that the
temporal cortex plays an important role in storing semantic information.
Semantic dementia is caused by degeneration of the neocortex of the
anterolateral temporal lobe (Lambon Ralph and Patterson, 2008). At least in the
early stages of the degenerative process, the hippocampal formation and the rest of
the medial temporal lobe are not affected.
 Reconsolidation of memory: Role of hippocampus
� What happens to memories of events as time goes on? If we learn something new about a particular subject,
our memories pertaining to that subject must somehow be modified.
� Maintenance of memory is an active process. Because a consolidated LTM can become susceptible to
disruption and restoration, a process termed “reconsolidation”.
� One of the side effects of a procedure known as electroconvulsive therapy is a period of retrograde amnesia.
The procedure, used to treat cases of severe depression, involves the application of electricity through
electrodes placed on a person’s scalp. The current excites so many neurons in the brain that it produces a
seizure. Presumably, the seizure erases short-term memories present at the time and thus prevents
consolidation of these memories.
� Misanin et al., (1968) found that long-term memories, which are normally not affected by seizures, were
vulnerable to disruption by electroconvulsive shock (ECS) if a reminder of the original learning experience
was first presented. The investigators found that ECS given right after a learning experience prevented
consolidation, but ECS given a day later did not. Apparently, the ECS given right after training disrupted the
brain activity initiated by the training session and consequently interfered with consolidation. The ECS given
the next day had no effect, because the memory had already been consolidated. However, if animals were
given a “reminder” stimulus one day after training, which presumably reactivated the memory, an ECS
treatment administered immediately afterward caused amnesia for the task when the animals were tested the
following day. Reactivation of the memory made it susceptible to disruption. These studies involved stimulus-
response learning.
� More recent studies have found that long-term, well-consolidated relational memories are also susceptible to
disruption. Presumably, the process of reconsolidation, which involves neural events similar to those
responsible for the original consolidation, makes it possible for established memories to be altered or
attached to new information (Nader, 2003).
� Events that interfere with consolidation also interfere with reconsolidation and can even erase memories or at
least make them inaccessible. For example, Debiec et al., (2010) trained rats on two fear conditioning tasks.
They paired two different tones (CSa and CSb) with shocks delivered to two different parts of the body (USa
and USb). After training, they randomly presented only one of the shocks (USa or USb) to each animal
followed by an infusion of anisomycin (which blocks protein production) or a placebo into the lateral
amygdala. Protein production is an important step in longterm potentiation and memory formation. The next
day, they tested the animals for retention of the conditioned responses. They found that the animals did not
freeze when they presented the tone that had been paired with the US that had been presented just before the
infusion of anisomycin on the previous day. In other words, presenting one of the shocks “reopened” the CS-
US memory for possible reconsolidation, which made the memory link fragile. Blocking protein synthesis
prevented the memory from going back to its original state.
 Hippocampal neurogenesis and consolidation
� Neurogenesis is the production of new neurons in the
brain
� Radial glia-like neural stem cells (RGLs) in
the dentate gyrus subregion of the hippocampus give
rise to dentate granule cells (DGCs) and astrocytes
throughout life, a process referred to as adult
hippocampal neurogenesis
� The newly generated neurons makes synaptic
connections with the dentate gyrus and CA 3.
� The rate of neurogenesis gradually declines with age,
however active
� The rate of neurogenesis and the survival of new
neurons are responsive to the changes in the
organism's environment.
� Antidepressant drugs increase hippocampal
neurogenesis whereas stress hormones decrease it.
 Morris (1982) water maze experiment on rats

� Neurogenesis in the
hippocampus in the Rats
exposed to relational
learning increased whereas
no effects on the rats
exposed to –R learning
� Because relational learning
involves hippocampus
whereas S-R learning does
not involve it.
 Role of cortex in semantic memory-Evidence

In semantic dementia, semantic information is lost, but episodic memory for

recent events can be spared.
The hippocampal formation and the limbic cortex of the medial temporal lobe appear
to be involved in the consolidation and retrieval of declarative memories, both
episodic and semantic, but the semantic memories themselves appear to be stored in
the neocortex—in particular, the anterolateral temporal lobe. Pobric et al., (2007)
found that TMS of the left anterior temporal lobe, which disrupts the normal neural
activity of this region, produced the symptoms of semantic dementia. Volunteers who
experienced TMS of the left anterior temporal lobe had difficulty naming pictures of
objects and understanding the meanings of words, but they had no trouble performing
other, nonsemantic tasks such as naming six-digit numbers and matching large
numbers according to their approximate size.
Surprise!!

1. Compare and contrast different types of learning providing

examples?
2. Explain the role of cerebellum and basal ganglia in motor
learning and memory?
3. The role of different regions of extra-striate cortex in visual
perceptual learning?