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Subject PSYCHOLOGY

Paper No and Title Paper No. 14: Neuropsychology

Module No and Title Module No. 30: Neural Mechanisms of Learning

Module Tag PSY_P14_ M30

TABLE OF CONTENTS
1. Learning Outcomes
2. Introduction
3. Neural Mechanisms of Classical Conditioning
3.1. Role of Amygdala
4. Operant Conditioning
4.1. Role of Basal Ganglia
5. Perceptual Learning
5.1. Role of Visual Cortex
6. Learning and Synaptic Plasticity: Role of Hippocampus
7. Chemical changes during Learning
8. Summary

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1. Learning Outcomes
After studying this module, you shall be able to;

 Learn about chemical changes that accompany learning Learn how learning may be
defined in a neurological paradigm
 Learn neural mechanisms of classical conditioning
 Learn neural mechanisms of operant conditioning
 Learn the neural mechanisms of perceptual learning
 Understand the concept and neurological basis of synaptic plasticity in learning

2. Introduction
The brain constantly rewires itself to become an organ that is physically different from what it
was in the preceding moment. Never resting, it churns through new experiences daily,
incorporating some new information into its network of stored information and discarding others.
The ever-changing nature of human brain gives rise to two questions that have been bothering
thinkers for centuries. They are;
 Do psychological processes end into biological processes?
 Can we physically locate all our experiences?

Aristotle’s teacher Plato reasoned that the mind had to exist inside the brain because of geometry
and pure logic. The brain was round he said, and close to the perfect roundness of the sphere. It
also inhabited the part of human body closest to heaven. Plato and other Greek philosophers
theorized about the existence of a force that keep people alive and left them at the time of death.
They called this force psyche or soul and several authors said it resided in the brain.

Centuries later, Galen, a Roman physician who lived in the eastern Mediterranean in the second
century of the Christian era, went beyond such mental exercises to test the brain himself. He took
a more hands-on approach and cut the sensory and motor fibers in pig’s brains to observe results.
Galen became the first to speculate that certain functions are carried out by specific parts in the
brain. He made rudimentary descriptions of the body’s major organs and fleshed out description
of what he saw as the varieties of human spirit. Thus was born the conundrum that has sparked
debate for many centuries. It is called the mind-body problem.

Neuropsychology has emerged as an important branch of Psychology to understand cognitive and

the higher order functions of organisms in the recent times. The phenomenon of learning,
memory, consciousness, self awareness, and perceptual processes has been the subject of analysis
in terms of their neurological basis. The biological and neurological substrate of various kinds of
learning has been delineated through scientific methods derived from physiology, biology,
chemistry and psychology. The mechanisms of learning have been studied in terms of braib
structures, as well as in terms of the bio-chemistry of the brain. Even molecular genetic studies
have been carried out to study learning processes in different paradigms.

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Definition of Learning in terms of Neurology

Learning and Memory make each individual unique. Even before birth, the human brain takes in
sensations, processes them and begins to encode them into trillions of synapses. Those
connections and the electrochemical firing patterns that unite them, make the brain an organ
different from what it was a moment before. As new connections form in response to stimuli and
then become strong through repeated use, the brain integrates new information and stores it until
needed. Without learning, memory would be little more than a clock work. Learning and memory
work together.
As a consequence of learning, an organism’s way of perceiving, thinking, feeling, and behaving
undergoes a change. The processes of learning produce changes in the neuronal synaptic
connections in various areas of the brain including those connected with motor behavior.
From a purely psychological point of view, learning may be defined as the relatively permanent
change in behavior acquired through practice and experience (Baron, 1998). Neural definition of
Learning, however, may be summarized as structural and chemical alterations that are acquired in
the brain on encountering an external stimulus.

3. Neural Mechanisms of Classical Conditioning

When Pavlov first described the process of Classical Conditioning, it was fairly behavioral in
nature, based on observable facts of the experiments. Later, as the belief of the role of neural
processes was understood, the neuropsychologists and cognitivists began to explain it in terms of
the neurological mechanisms involved. Neuroscientists have studied the anatomy and physiology
of classical conditioning using many models such as the Gill Withdrawal Reflex in Aplysia
(Levond et al, 1993).

In classical conditioning, the stimulus (S) triggers the response (R) of an organism. With the
exposure of the organism to the stimulus, reflex results. Reflex is the involuntary behavior which
comes from within. One of the most common reflexes seen is the blinking of the eye whenever air
is blown into it or a foreign object approaches it. The concept of reflex, with no conscious
control, is incorporated in classical conditioning.

Pavlov developed a model for classical conditioning. It involves the following;

 The Unconditioned Stimulus (US) triggers the Unconditioned Response (UR). This
means that without learning, a stimulus can elicit a reflex. Reflex is called the
unconditioned response because it is involuntary and we do not need to learn it for the
event to occur.
 The Neutral Stimulus (NS) is a stimulus that does not elicit the Unconditioned Response
(UR). This means that when this stimulus is presented to the organism, it would not
execute the same response he showed when it encountered US.
 US and NS are repeatedly paired and presented to the organism in conjunction
 The pairing of US and NS transforms NS into a Conditioned Stimulus (CS). This means
that whenever the previously neutral stimulus is presented alone (US is not showed
anymore) to the organism, it causes UR to occur. But this time, UR is changed into
Conditioned Response (CR), because the response is elicited due to conditioning.
Therefore, CS elicits CR.

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Classical Conditioning

Pavlov believed that classical conditioning reflected a strengthened connection between a brain
area that represents CS activity and a brain area that represents UCS activity. Because of that
strengthened connection, any excitation of the CS centre flows to the UCS center evoking the
unconditional response. The connection between these two areas was labeled as an engram.
Lashley (1950) studied such engrams by severing the connection between several brain areas
after training subjects in classical conditioning. No specific place where such engrams are made
was discovered. Learning was found to take place in diffuse areas.

Analysis of classical conditioning was presented in the form of a model by Weinberger and his
colleagues. The paradigm used was that of the conditioning of the emotional response. The
unconditioned stimulus was the food, and the conditioned stimulus was the sound, a tone of a
certain frequency. In the presence of unconditioned stimulus, i.e. food, not only salivation but
also the autonomic arousal took place. The medulla hormone, as well as the neurotransmitter nor
epinephrine was also released. With repeated pairings, the conditioned stimulus of tone resulted
in similar release of the neurotransmitter. Nor epinephrine increases the level of glucose in blood
which facilitates the process of learning and memory.

3.1. Role of Amygdala

At the level of neurological structures, the response to food travels via a sensory pathway that
runs through the central nucleus of the amygdala to the somato-sensory cortex. This is not
surprising, given that the central nucleus plays an important role in emotion. The auditory
stimulus, the tone of a given frequency, travels via a sensory pathway that passes through the

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medial geniculate nucleus of the thalamus and on to the

auditory cortex. Pathways of the two are diagrammatically
represented below:

CS: Lateral nucleus amygdala

via thalamus
UCS: Somatosensory cortex via
central nucleus of amygdala

CR

Amygdala is an important structure located within the depth of the temporal cortex, and is a
significant component of the limbic system of the brain. The limbic system, with its circular
circuitry, is the major center for emotions. It has a big role in organizing the pattern of emotional
responses evoked by the unconditioned stimulus. Central nucleus of Amygdala is the specific site
where activation is observed at the time of emotional experience. The information about
conditioned stimulus reaches at the lateral nucleus of amygdale and that about the unconditioned
one from the somato-sensory system. These two sources of information converge in the lateral
nucleus resulting in learning of the Conditioned Response.

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Figure showing Association of auditory and somato-sensory information

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Besides amygdale, the role of the nucleus basdalis has also

been highlighted. It is shown as the place where junction
between somato sensory messages and the information coming from the auditory cortex. The
nucleus basalis contains neurons having acetylcholine as the neurotransmitter, and have a
diffused system of projection to the cerebral cortex. Such diffused activation also increases the
sensitivity of the auditory cortex to receive signals activation of the cerebral cortex also increases
the sensitivity of the auditory cortex to receive information (tone) from the medial geniculate
nucleus. Thus, it is via the nucleus basalis, acetylcholine containing neurons, and the auditory
cortex, that the unconditioned stimulus (the food) and conditioned stimulus (the tone) get
associated.

4. Operant Conditioning
Instrumental or operant conditioning is the means by which we profit from experience. If in a
particular situation, we make a response that had favorable outcomes we tend to make the
response again.

4.1. Role of Basal Ganglia

Operant Conditioning entails the strengthening of connections between neural circuits that detect
a particular stimulus and neural circuits that produce particular response. Therefore, instrumental
conditioning involves various regions of sensory association cortex where the perception takes
place and the motor association cortex of the frontal lobe.
There are two major pathways between the sensory association cortex and the motor association
cortex-
 Direct trans-cortical connections
 Connections via basal ganglia and thalamus
Both of these pathways seem to have a crucial role in Operant Conditioning. However they
function differently.
The direct trans-cortical connections are involved in the acquisition of episodic memory and in
acquisition of complex behaviors such as following instructions. It helps establishing rules (this
leads to that) and thus, in future, in case of similar situation that present similar stimuli, these
connections offer information as learnt earlier.
Evidence suggests that as learned behaviors become routine or automated they are transferred to
basal ganglia. The basal ganglia receive information about the stimuli that are present and the
responses that we are making. When these are repeated, it is stored here. And once again, the
trans-cortical connections are free for other tasks. The outputs of the basal ganglia are sent to the

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frontal cortex, to the pre-motor and supplementary motor

cortex where the plans of movements are made. Finally, it travels to primary motor cortex where
the plans of actions are executed.
.

Activation of basal ganglia during operant conditioning

5. Perceptual Learning
Learning involves adaptation to the changing environment and to coordinate behavior
accordingly. The behavior constantly undergoes changes as the situation demands. Getting to
understand the environment and the changes occurring in it, the kind of learning required is called
the perceptual learning. It is a life-long learning process si9nce the situations keep changing. It
requires encoding and assimilation of information.

Perceptual learning has a wide neurological basis. It is largely located in the temporal cortex of
the cerebral cortex. Attempts to find out neural base of prospagnosia, a failure to recognize faces,
led to the discovery that the neural substrate for perceptual learning are largely located in the
temporal cortex. Lesions in this area produced disruption in this kind of learning.

5.1. Role of Visual Cortex

When the unconditioned stimulus is visual in nature, the visual cortex is actively involved.
Objects are recognized visually by circuits of neurons in the visual association cortex. The visual
cortex receives information from the lateral geniculate nucleus of the thalamus. Within the
primary visual cortex, individual modules of neurons analyze the information from restricted
regions of visual scene. The information is collected in the sub-regions of extra-striate which
surround the primary visual cortex. The ventral stream of the visual cortex is associated with the
object recognition. The dorsal stream is involved in the perception of location.

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6. Learning and Synaptic Plasticity: Role of Hippocampus

Since learning is an ongoing process, in needs to be based on neural structures having flexibility
or plasticity in their structure and function. It is required at the bio chemical level where synaptic
connections between neurons take place. They may later be represented in terms of brain
structures also..
When re-learning of some structures is involved, hippocampus takes up a big role. Synapses have
plasticity and small alterations in the learning structures go on happening.
Long term potentiation is essential for the consolidation of learning. When an input (learning)
produces activation repeatedly, it leads to an increase in the excitability of these neurons on a
long term basis. This is called the process of long term potentiation. The evidence for such a
potentiation has been observed in the hippocampal areas of the brain. The hippocampus is also
located in the temporal cortex and is an important part of the limbic system. When high frequency
stimulation of axons in hippocampus is administered experimentally, it leads to the strengthening
of synaptic connections. It is found to increase the size of excitatory post synaptic potentials in
the post synaptic neurons. Even weak synapses may be strengthened by the action of strong ones.
This process is known as the associative long term potentiation.
Hippocampal formation is a fore brain structure of the temporal lobe, constituting an important
part of the limbic system, and includes dentate gyrus and subiculum. CA1 and CA3 are important
structures of the hippocampal formation.
In the field CA1 and dentate gyrus. NMDA receptors play special role in long term potentiation.
NDMA receptors are specialized ionotropic glutamate receptors that control the calcium
channels. These receptors respond to glutamate, and control calcium channels. However, they can
open them only when the membrane has been depolarized. Calcium ions can enter into the cells

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only when depolarization of the receiving membrane and

activation of the NMDA receptors is taki8ng place
simultaneously. Increased level of calcium in the cell activates other enzymes to allow AMPA
receptors to get activated. The AMPA is a receptor containing glutamate and can control the
sodium channels. It is, thus, involved with producing excitatory post synaptic potentials. These
changes result in structural alterations in the shape of dendrites. Long term potentiation requires
protein synthesis which appears to take place in the dendrite adjacent to dendritic spine.
Another interesting finding is that instead of long term potentiation, the long term depression can
also occur. It happens when the synapse occurs coinciding with the hyper polarization of the
receiving post synaptic membrane.
The process of Long term potentiation may be summarized below;
After repetitive stimulation of a neural pathway, a synapse will show two signs of increased
efficiency:
1. A single stimulation causing a more rapid increase in the Excitatory Post Synaptic Potential
(EPSP).

2. A single stimulation recruiting more cells that reach the action potential threshold.

7. Chemical Changes during Learning

1. Synaptic activity in hippocampus releases Glutamate (excitatory) and Calcium ( Ca++ )

 NMDA (a type of receptor) is blocked by Mg++

 AMPA (a type of receptor) opens

2. AMPA allows in Sodium (Na+)

3. Excitatory Postsynaptic Potential (EPSP) results

4. Activation of postsynaptic dendrite releases Mg++ from NMDA receptors

5. Glutamate binds with NMDA receptor

6. NMDA pumps Ca++ into cell and causes Action Potential faster than Na+

7. Ca++ used to synthesize neuro-trophins.

8. Neurotrophins affect synapse (more, larger, stronger connections)

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Long term potentiation causes AMPA receptors to move into the dendritic spines and appears to
cause the formation of new synapses. Chemical reactions not yet fully understood also cause the
release of glutamate. Long term potentiation also requires synthesis of new proteins including
components of cytoskeleton, protein kinesis, and receptors. It is also accomplished here in this
process.

8. Summary
 Learning accompanies structural and chemical alterations in the brain on encountering an
external stimulus.
 The central and lateral nucleus of amygdala play significant role in the process of
classical conditioning.
 Direct trans-cortical connections and those via basal ganglia and thalamus are involved in operant
conditioning
 Visual cortex plays important role in perceptual learning
 Long term potentiation or consolidation of learning takes place in the hippocampal
formation where NMDA and AMPA receptors facilitate the process via their chemical
mediation.

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