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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr.

Nwogueze BC

PIO203
Neurophysiology I (Introduction to
Neurophysiology)

By

Dr. Nwogueze BC

OUTLINE
1. Introduction to neurophysiology
2. Neurons
3. Classification of nerve fibres, properties of nerve fibres, neuroglia
4. Receptors
5. Synapse
6. Neurotransmitters
7. Reflex activity and reflex arc
8. Spinal cord

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

INTRODUCTION TO NEUROPHYSIOLOGY
The common approach of studying Physiology is system wise. System is a group of organs and
structures concerned with a common body functions.
1. Circulatory system or Cardiovascular system
2. Respiratory system
3. Excretory system
4. Digestive system
5. Endocrine system
6. Nervous system
7. Special senses
8. Musculo-skeletal system
9. Reproductive system

Many functions of living organisms are regulatory processes- a certain state, characterized by a
measurable quantity, is kept constant. Examples are the regulation of body temperature, of blood
pressure, and of the position of the body in the gravitational field. In many cases of regulation,
the nervous system plays a central role. Each system is composed of a set of inter connected and
interdependent organs which act together for a common purpose.

Nervous System
Nervous system is a complex and well-organized system of communication that allows
the individual to interact with his external environment. It is also a system of integration that
coordinates functions of various internal organ systems of the body. Neural connections between
body parts provide the basis for anatomical and physiological communications that help in
smooth execution of most of the systemic functions such as gastrointestinal functions, secretion
of hormones, functions of heart, lungs and kidney, and musculoskeletal system etc

One easy way to begin to understand the structure o f the nervous system is to start with the
large divisions and work through to a more in-depth understanding.

The nervous system consists of two broad divisions: Central nervous system (CNS) and
peripheral nervous system (PNS).

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

a. Central Nervous System


The central nervous system (CNS) includes brain and spinal cord. Brain is situated within the
cranial cavity of the skull, and it continues into the vertebral canal as the spinal cord, i.e the
spinal cord is contained within the vertebral cavity of the vertebral column.

1. Brain: The brain is divided into three parts namely;


i-Prosencephalon i.e the forebrain: consists of the telencephalon (the two cerebral
hemispheres) serves for perception of sensations, cognition, learning and memory, and planning
and programming of responses and the diencephalon (thalamus, hypothalamus, metathalamus,
and subthalamus) for relay of information to cortex, and control of autonomic and endocrine
functions.
ii-Mesencephalon i.e the midbrain: consists of the corpora quadrigemina, cerebral peduncles,
substantia nigra, tegmentum and many midbrain nuclei for locomotion and nuclei for righting
reflexes. It also contains a part of reticular formation.

iii-Rhombencephalon i.e the hindbrain: consists of metencephalon (the pons and the
cerebellum) and myelencephalon (the medulla oblongata).

NB: The midbrain, pons, and medulla forms the brainstem. The main functions of brainstem are
control of cardiovascular and respiratory functions, motor activities, sleep-wakefulness and
visceral functions.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

Diagram showing Parts of CNS


b. Peripheral Nervous System
The peripheral nervous system (PNS) includes somatic nerves that come out of the spinal cord
to innervate different body structures like skin, muscles, bones, etc. The structures of the PNS
are referred to as ganglia and nerves, which can be seen as distinct structures. The main function
of PNS is to collect and convey information from peripheral structures to CNS via sensory
axons and convey motor signals from CNS to muscles and related structures via motor neurons.

The nervous system can be divided into two parts mostly on the basis of a functional
difference in responses.
a. somatic nervous system (SNS)
The somatic nervous system (SNS) is responsible for conscious perception and
voluntary motor responses. Voluntary motor response means the contraction of skeletal
muscle, but those contractions are not always voluntary in the sense that you have to want
to perform them. Some somatic motor responses are reflexes, and often happen without a
conscious decision to perform them. If your friend jumps out from behind a corner and

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

yells “Boo!” you will be startled and you might scream or leap back. You didn’t decide to
do that, and you may not have wanted to give your friend a reason to laugh at your
expense, but it is a reflex involving skeletal muscle contractions. Other motor responses
become automatic (in other words, unconscious) as a person learns motor skills (referred
to as “habit learning” or “procedural memory”).
b. autonomic nervous system (ANS)
The autonomic nervous system (ANS) is responsible for involuntary control of the
body, usually for the sake of homeostasis (regulation of the internal environment).
Sensory input for autonomic functions can be from sensory structures tuned to external or
internal environmental stimuli. The motor output extends to smooth and cardiac muscle
as well as glandular tissue. The role of the autonomic system is to regulate the organ
systems of the body, which usually means to control homeostasis. Sweat glands, for
example, are controlled by the autonomic system. When you are hot, sweating helps cool
your body down. That is a homeostatic mechanism. But when you are nervous, you might
start sweating also. That is not homeostatic, it is the physiological response to an
emotional state.
Functions of The Nervous System
The major functions of the nervous system are:
1. Communication between body parts and integration of systemic functions.
2. Appropriate regulation of body functions.
3. Coordinated interaction of the body with the environment.
The neural regulation is the major controlling mechanism of many functions and processes of
the body. Nervous system achieves its objectives through neurons that are designed for rapid
transmission of information from one body part to the other.

CNS accomplishes its functions through the following major processes:


i. Recognition of stimulus and transmission of information in the form of nerve impulse
to CNS via sensation (changes in the form physical energy from the environment),
receptors (transduce environmental energy into the action potentials; NB- forms of
stimuli that are detected by receptors include mechanical, chemical, photic (light),
auditory (sound), thermal (temperature), and electrical), Sensory neurons (transmit

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

information in the form of nerve action potentials from receptors to the CNS via
sensory pathways, NB refer to special senses)
ii. Information Processing and Generation of Command Signal via action potential
((sensory signal) transmitted is first relayed in the thalamus and then processed in
sensory cortex, thus, the processed signals are transformed into sensory networks in
the brain); Integration of the processed signals into appropriate command signals
and then Transmission of the command signal to the effector organs for
implementation of the plan. Then, learning based on sensory inputs and then storage
of learned information in memory for future utilization of the knowledge
iii. Specific responses in accordance to the command signal conveyed by CNS via
movement of body parts (motor activities), change in visceral functions (autonomic
responses) or even the change in behavior of the individual.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

NEURONS
Definition:

The building blocks of the nervous system are the nerve cells, also called neurons. A neuron is
the structural and functional unit of the nervous system. Its primary function is to receive the
various stimuli and transmit the signals to other neurons and tissues (pain, touch, light, sound,
cold and warmth). Neurons have evolved from primitive neuroeffector cells that respond to
various stimuli by contracting. It is estimated that the human brain possesses 25 billion cells. In
complex animal, integration and transmission of nerve impulses have become the specialized
functions of neurons (action potentials, receptor potentials, and synaptic potentials).

Classification:
The classification of neurons can be on basis of; the length of the axon, number of poles,
dendritic pattern and their functions.
1. Based on the length of Axon
Neurons can be classified into 2 based on the length of axon
i. Golgi Type I: These types of neurons have long axons. Cell body of these neurons is in
different parts of central nervous system and their axons reach the remote peripheral organs. The
example is cortical inhibitory neurons.
ii. Golgi Type II: These types of neurons have short axons. These neurons are present in
cerebral cortex and spinal cord. Cortical motor neurons (neurons that give rise to corticospinal
tract) are the examples.
2. Based on the number of Poles
Neurons can be classified into 4 based on the number of poles;
i. Unipolar Neurons: Neurons that have only one pole are called unipolar neurons. From a
single pole, both axon and dendrite arise. This type of nerve cells is present only in embryonic
stage in human beings.

ii. Pseudounipolar Neurons: In this type of neurons, axon after originating from soma splits
into central and peripheral processes. The example is dorsal root ganglion cell (primary
sensory neurons with cell bodies in dorsal root ganglion).

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

iii. Bipolar Neurons: Neurons with two poles are called bipolar neurons. Axon arises from one
pole and dendrites arise from the other pole.

iv. Multipolar Neurons: Neurons which have many poles are called multipolar neurons. One
of the poles gives rise to axon and all other poles give rise to dendrites.

Figure 2: Types of Neurons

3. Based on the dendritic pattern of neurons:


Neurons can be classified into 2 based on dendritic pattern;
i. Pyramidal Cells: Dendrites of these cells spread like pyramids. The example is hippocampal
pyramidal neurons.

ii. Stellate Cells: Radial shaped spread of dendrites occurs in these cells. The examples are
cortical stellate cells.

4. Based on the functions of neurons:


Neurons can be classified into 2 based on functions;
i. Sensory or Afferent Neurons: These are neurons which carry the sensory impulses or
sensation from periphery to central nervous system. Generally, each sensory neuron has a short
axon and long dendrites.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

ii. Motor or Efferent Neurons: These are neurons which carry the motor impulses from central
nervous system to peripheral effector organs like muscles, glands, organs, system, blood vessels,
etc. Generally, each motor neuron has a long axon and short dendrites.

Structure of Neuron:
The size and shape of neurons vary widely, but the structural plan always includes 3 main
elements namely: a nerve cell body or soma, and the processes from this cell body, namely an
axon (neurite), and usually several dendrites.

Figure 1: Schematic diagram of a neuron showing the soma, the axonal and dendritic processes,
and the cell contents

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

Figure 2: Detailed structure of the proximal part of neuron,

a. Nerve Cell Body


The nerve cell body of soma or perikaryon is irregular in shape. Like any other cell, it is
constituted by a mass of cytoplasm called neuroplasm, which is bounded by a cell
membrane. The plasma membrane of the soma is a bilayered lipid protein membrane,
called plasmalemma. The cytoplasm contains a large nucleus, Nissl bodies, neurofibrils,
mitochondria and Golgi apparatus. Nissl bodies and neurofibrils are found only in nerve
cell and not in other cells. The nerve cell body helps to maintain the functional and
anatomical integrity of the axon. The proteins associated with synaptic transmitters are
synthesized in Nissl granules of the cell body and are transported to axon terminal by
axoplasmic flow.
i. Nucleus: Each neuron has one nucleus, which is centrally placed in the nerve cell
body. Nucleus has one or two prominent nucleoli. Nucleus does not contain
centrosome, thus, nerve cell cannot multiply like other cells.
ii. Neurofibrils: this is also called the neurofilaments with features of thread-like
structures/networks present in the form of network in the soma and the nerve
processes. The neurofibrils consist of microfilaments and microtubules.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

iii. Mitochondria: These organelles are present in soma and in axon. As in other cells,
here also mitochondria form the powerhouse of the nerve cell, where ATP is
produced.
iv. Nissl bodies: this is also called the Nissl granules are small basophilic granules or
stacks of rough endoplasmic reticulum found in cytoplasm of neurons. These bodies
are present in soma and dendrite b+ organelles containing ribosomes and polysomes.
So, these bodies are concerned with synthesis of proteins in the neurons. Proteins
formed in soma are transported to the axon by axonal flow. Number of Nissl bodies
varies with the condition of the nerve.
v. Golgi apparatus: The Golgi apparatus of nerve cell body is similar to that of other
cells. It is concerned with processing and packing of proteins into granules.

b. Dendrite
Dendrite is the branched process of neuron and it is branched repeatedly. This implies
that neurons have several processes called dendrites that extend outward from the cell. In
the cerebral and cerebellar cortex, the dendrites have small knobby projections called
dendritic spines. Dendrite may be present or absent. If present, it may be one or many in
number. Dendrite has Nissl granules and neurofibrils. Dendrite receives the incoming
signals from other cells and transmits the impulses towards the nerve cell body. Usually,
the dendrite is shorter than axon. On an average, there are about 10,000 dendrites per
neuron present in the CNS. Dendrites form the receptor zone of the neuron, i.e. they
receive impulses and transmit the impulses toward the cell body. In this region, non-
conducted local potential changes generated by synaptic connections are integrated.
c. Axon
Axon is the longer process of nerve cell. Each neuron has only one axon. Axon arises
from axon hillock of the nerve cell body and it is devoid of Nissl granules. Axon extends
for a long distance away from the nerve cell body. Length of longest axon is about 1
meter. Axon is a long tubular process that extends away from the nerve cell body to
transmit output signals to target organs. The initial segment of the axon is the site where
propagated action potentials are generated. The axonal process transmits propagated
impulses from the cell body to the axon terminal.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

Organization of Nerve
Each nerve is formed by many bundles or groups of nerve fibers. Each bundle of nerve
fibers is called a fasciculus. The whole nerve is covered by tubular sheath, which is formed by
areolar membrane. This sheath is called epineurium. Each fasciculus is covered by
perineurium and each nerve fiber (axon) is covered by endoneurium.

Figure 3: Organization of a nerve

Internal Structure of Axon


An axon has a long central core of cytoplasm called axoplasm. The axoplasm is covered
by the tubular-like sheath membrane called axolemma which is the continuation of the cell
membrane of nerve cell body. The combination of axoplasm with axolemma forms the axis
cylinder of the nerve fiber which is covered by a membrane called neurilemma. Axoplasm
contains mitochondria, neurofibrils and axoplasmic vesicles. Because of the absence of Nissl
bodies in the axon, proteins necessary for the nerve fibers are synthesized in the soma and not in
axoplasm. After synthesis, the protein molecules are transported from soma to axon, by means of
axonal flow. Some neurotransmitter substances are also transported by axonal flow from soma to
axon.

Types of Axon: Myelinated and Non-myelinated Nerve Fiber):

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

The non-myelinated nerve fiber is not covered by myelin sheath while nerve fiber which
is insulated by myelin sheath is called myelinated nerve fibers. Myelin sheath is a thick
lipoprotein sheath is formed by the Schwann cells that insulate the myelinated nerve fiber.
Myelin sheath is not a continuous sheath. It is absent at regular intervals. The area where myelin
sheath is absent is called node of Ranvier. Segment of the nerve fiber between two nodes is
called internode. Myelin sheath is responsible for white color of nerve fibers.

Figure 4: Myelinated nerve fiber (A); Non-myelinated nerve fiber (B).

Figure 5: Myelin sheath, shown in transverse section (A) and longitudinal section (B) of the axon

Functions of Myelin Sheath


i. Myelin sheath has a high insulating capacity. Because of this quality, myelin sheath
restricts the nerve impulse within single nerve fiber and prevents the stimulation of
neighboring nerve fibers.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

ii. Myelin sheath is responsible for faster conduction of impulse through the nerve
fibers. In myelinated nerve fibers, the impulses jump from one node to another node.
This type of transmission of impulses is called saltatory conduction.
iii. It is responsible for the color of the white matter of the brain and spinal cord.
iv. It reduces energy expenditure by the cell.

Axoplasmic Transport
Transfer of substances between cell body and axon terminal is called axoplasmic transport.
Various proteins, organelles and other cellular substances required for the development, growth,
and maintenance of the neuron are transported mainly along the length of the axon. In the
axoplasm, transport process can occur in both directions by different transport mechanisms.
Accordingly, they are called anterograde, retrograde, and transneuronal transports as described.
i. Anterograde or Orthograde transport occurs along microtubules that run along the
length of the axon and requires two molecular motors; dynein and kinesin.
Orthograde transport moves from the cell body toward the axon terminals. For
example, various neurotransmitters synthesized in the cell body are packaged in
vesicles and get secreted at the nerve endings through axoplasmic microtubules. The
rate of transport process may be fast or slow;
 Fast axonal transport occurs at about 400 mm/day (which is accomplished by
kinesin, a microtubule associated protein that transports many organelles, vesicles
and membrane glycoproteins)
 Slow axonal transport occurs at 0.5 to 10 mm/day (Various structural proteins
like actin, neurofilaments and microtubules get transported by slow transport).
ii. Retrograde transport this involves the transport of substances from the axon
terminals to the cell body is known as retrograde transport. It occurs at a speed of
about 200 mm/day, brought about by dynein, another microtubule associated protein.
This mechanism keeps the soma informed about the synaptic environment. The
examples of retrograde transport are:
 Transport of toxins: Tetanus toxin at motor neuron ending is transported to the
cell body by this retrograde process.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

 Transport of viruses: The chickenpox virus, known as varicella zoster that


causes herpes simplex reaches cell body from nerve terminals in the skin by
retrograde transport.
 Reuptake of synaptic transmitters: Neurotransmitters like norepinephrine (NE)
released at the nerve terminals are rapidly removed from the synaptic cleft by
reuptake into the presynaptic neuron.
iii. Transneuronal Transport involves the transport of trophic substances like nerve
growth factors across the synapse to the presynaptic membrane of another neuron.
This is called transneuronal transport. This helps in maintenance of the synaptic
contacts.
Neurotrophins (Neurotrophic Factors)
Neurotrophins are secreted by many tissues in the body, particularly muscles, neuroglial
cells called astrocytes and neurons. Neurotrophins act via neurotrophin receptorsbinding, which
are situated at the nerve terminals and nerve cell body. Neurotrophins or neurotrophic factors are
the protein substances, which play an important role in growth and functioning of nervous tissue
(CNS and PNS), they promote survival and repair of the nerve cells and play an important role in
the maintenance of nervous tissue and neural transmission.
Types of Neurotropins
i. Nerve Growth Factor (NGF): promotes early growth and development of neurons
ii. Ciliary Neurotrophic Factor (CNTF): protects neurons of ciliary ganglion and
motor neurons
iii. Brain-derived neurotrophic growth factor (BDGF): promotes the survival of
sensory and motor neurons
iv. Neurotrophin-3 (NT-3): regulates the release of neurotransmitter from
neuromuscular junction
v. Neurotrophins 4 and 5: they act through tyrosine kinase B. Exact function is not
known.
vi. Glial Cell Line-derived Neurotrophic Factor (GNDF): potent protective action on
dopaminergic neurons
vii. Fibroblast Growth Factor (FGF): promotes fibroblastic growth and protect the
neurons.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

CLASSIFICATION OF NERVE FIBRES, PROPERTIES OF NERVE


FIBRES, NEUROGLIA
1. CLASSIFICATION OF NERVE FIBERS
There are many classifications of nerve fibers, broadly there are 2 Classification systems of
Nerve Fibers, namely Erlanger and Gasser system (All the nerve fibers i.e. sensory, motor and
autonomic fibers) and the Lloyd system (sensory nerve fibers)

i. Erlanger and Gasser


The Erlanger and Gasser is the most popular classification of nerve fibers, based on their
diameter and conduction velocities. Broadly fibers are classified into three categories: A, B and
C.

 Type A Nerve Fibers


These are the fastest conducting fibers that are myelinated with a conduction velocity ranging
from 70 to 120 m/sec. The fiber diameter varies from 12 to 20μm. Type A fibers are further
subdivided into α, β , γ and δ fibers.
1. Aα fibers (somatic sensory and somatic motor) supply extrafusal fibers in skeletal muscles
and are for proprioception.
2. A β fibers (sensory) carries vibration, proprioception, and light touch sensations.
3. Aγ fibers (motor) supply intrafusal fibers in muscle spindles.
4. Aδ fibers (sensory) are mainly fast pain and cold temperature conducting nociceptive fibers.

 Type B Nerve Fibers

These are preganglionic autonomic fibers (motor) that are myelinated. They have a diameter
of less than 3μm and conduction velocity ranging from 3 to 15m/sec.

 Type C Nerve Fibers


These are unmyelinated fibers (sensory). They are subdivided into two broad categories: dorsal
root fibers have diameter of 0.4 to 1.2μm and their conduction velocity varies from 0.5 to 2
m/sec, and postganglionic autonomic fibers (sympathetic) with diameter 0.3 to 1.3μm and
conduction velocity of 0.7 to 2.3 m/s. They C fibers carry various sensations (vibration, slow
pain, crude touch, temperature [cold and warm]

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

2. Lloyd (Numerical) Classification


These are the sensory nerve fibers and are typed into IA, IB, II, III and IV.
Types Origin Fiber types
IA Primary muscle spindle afferent from annulospiral ending Aα
(Senses changes in Length)
IB Golgi tendon organ Aα
(Senses changes in Tension)
II Secondary muscle spindle afferent from flower spray endings
Also from touch and pressure receptors Aβ
III Mainly from pain and cold receptors Aδ
IV Mainly from pain and temperature C fibers in
dorsal root
2. PROPERTIES OF NERVE FIBERS
a. Excitability
Excitability is the property by virtue of which cells or tissues respond to changes in the external
or internal environments. It is due to the disturbances in the ionic equilibrium across the
receptive zone of cell membrane. The nerve fibers are highly excitable tissues (ability to generate
electrical impulses or action potential). Nerve fibers respond to various forms of stimuli namely;
mechanical, thermal, chemical or electrical. In experiment set-up, ‘electrical’ stimulus is usually
employed, because its strength and frequency can be accurately controlled (easily be quantified),
least damaging to the nerve fiber, the phenomenon of excitability and conduction produced by
stimulus is same as natural stimulus, also nerves respond well to chemical and thermal stimuli.
There are 5 types of stimulus depending on strength namely; supramaximal, maximal,
submaximal, threshold/minimal, subthreshold stimulus. Factors Affecting Excitability includes;
strength and duration of the stimulus, as well as ionic concentration/distribution across the
membrane.

b. Conductivity
On stimulation, action potential is generated in the nerve fiber, which is propagated along its
entire length to the axon terminal. An axon can conduct in either direction. If the stimulus is
applied in the middle junction of axon, the action potential initiated in the middle of it can travel

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

in both directions, due to set-up of electronic depolarization on either side of the initial current
sink. Impulses normally pass from synaptic junction to the axon terminal, which is called
orhtodromic conduction. Conduction in the opposite direction is called antidromic conduction,
seen in sensory nerve supplying the blood vessels. Axon reflex is an example of antidromic
conduction. Factors affecting conduction of nerve fibers; temperature, resting membrane
potential, firing level, axon diameter, myelination, and resistance offered of ECF.

c. Unfatigability
Nerve fibers cannot be fatigued, even when they are stimulated continuously. This is because the
nerve fibers primarily conduct impulses (propagation of action potential) that do not involve
expenditure of energy (ATP).

d. Accommodation
Application of continuous stimuli may decrease the excitability of the nerve fiber, a phenomenon
called accommodation. More than nerve fiber, it is nerve endings that adapts. This decreases the
transmission of impulse across the neurons. If a nerve is submitted to the passage of constant
strength of current, the site of stimulation shows decrease in excitability. The accommodation
consists of a rise in threshold of the membrane during stimulation. A similar feature observed at
nerve endings is called adaptation. Thus, nerve fiber accommodates while the nerve endings
adapt.

e. Summation
Application of a subthreshold stimulus does not evoke an action potential. However, if
subthreshold stimuli are applied in rapid succession, they are summated and they produce an
action potential. This property is called summation.

f. Refractory Periods
During the action potential, the stimulated area of the membrane happens to be unresponsive to a
second stimulus in most part, and later it requires a stronger stimulus to get excited again. The
length of time during which the membrane is unresponsive to a second stimulus no matter

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

how strong is the stimulus, is known as refractory period. The periods of total and relative
refractoriness are known as absolute and relative refractory periods respectively.

Fig. 1: Refractory periods of nerve action potential. (ARP: Absolute refractory period; RRP:
Relative refractory period).

i. Absolute Refractory Period (ARP) is defined as the period in the action potential
during which, application of a second stimulus of any strength and duration does not
produce another action potential. The ARP corresponds to the period from the time
the firing level is reached until repolarization is about one-third complete. At the peak
of the action potential, the inactivation gates of the voltage-gated sodium channels
close and they remain in that inactivated state for some time before returning to the
resting state. These sodium channels can reopen in response to a second stimulus,
only after attaining the resting state. Hence, even if a stronger stimulus is applied
during this interval, it will not produce a second action potential, and the membrane is
said to be in its absolute refractory period.
ii. Relative Refractory Period (RRP) is defined as the period following ARP during
which, application of a suprathreshold stimulus can elicit a second action potential.
The RRP starts from the end of ARP to the start of after-depolarization. The
following factors contribute to RRP: All the sodium channels present at the site of
stimulus do not achieve the open state or inactivated state or resting state, exactly at

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

the same time. Few of them open when the membrane potential is –63 mV, causing
local response. By the time of relative refractory period, some of the channels have
returned to their initial resting state. These channels in resting state can open their
activation gate and allow the influx of Na+. Similarly, a suprathreshold stimulus can
spread to larger area over the membrane and open extra voltage-gated sodium
channel. Thus, only a suprathreshold stimulus and not any threshold stimulus can
open up sufficient number of sodium channels to elicit an action potential during the
RRP.

g. All-or-None Law
All or None Law states that the action potential occurs with a constant amplitude and shape
irrespective of magnitude of the stimulus. A subthreshold stimulus fails to excite the tissue.
Only a stimulus of threshold magnitude elicits an action potential. If suprathreshold stimuli
are applied, the action potentials resulting from them have the same amplitude, duration and
form as those produced by threshold stimuli, provided the experimental conditions like electrical
potentials on both sides of the membrane, concentration of ions in ICF and ECF, temperature’
etc. remain same. The activation gates of voltage-gated Na+ channels open as soon as 15 mV of
depolarization is achieved. Following that, any extra degree of depolarization is of no further
use, as the membrane automatically achieves another +90mV of depolarization (–55 to +35 mV):
i. Thus, after the threshold level is achieved, the amount of sodium influx becomes
independent of the stimulus factor.
ii. The number of voltage-gated Na+ channels over the axonal membrane of
unmyelinated axons as well as at the nodes of Ranvier in myelinated axons remains
fairly constant.
iii. Once the action potential is formed, it appears with its maximum size and shape,
otherwise it does not form at all.
iv. Therefore, the action potential follows the all-or-none law; i.e. the action potential
occurs with a constant amplitude and shape whether the stimulus is of threshold or
suprathreshold magnitude.

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PIO203- Neurophysiology I (Introduction to Neurophysiology) By Dr. Nwogueze BC

Fig 2: All or none law for action potential. Note, sub-threshold stimuli (1 and 2) do not evoke
formation of action potential, whereas, threshold (3and 4) and supra-threshold (5 and 6) stimuli
evoke similar action potentials.

3. NEUROGLIA
The nervous system aside the neurons contains several types of supporting cells called neuroglia.
Glial cells of nervous system are called neuroglia. Neuroglias support and protect the neurons
and maintain homeostasis of fluids that bath the neurons. Neurons transmit impulses in the form
of action potentials. Neuroglias are 10 – 30 times plentiful than neurons. Glial cells neither
conduct action potential nor form functional synapse with other cells. However, they can be
passively polarized in response to nearby neural activity. Though glial cells generally provide
support for neurons, their functions are complex and not completely understood.

Types of Glial Cells


Four types of glial cells have been described in CNS: astrocytes (astroglia), oligodendrocytes
(oligodendroglia), ependymal cells and microglia. Schwann cells are glial cells in PNS.

a. Astrocytes
Astrocytes are so named because of their star shape. They are found abundantly throughout the
brain and spinal cord. Processes of astrocytes surround the neurons and their axons, and often
terminate on the wall of blood vessels. Functions of astrocytes are as follows: they provide the
mechanical matrix, they serve metabolic and nutritive functions for neurons, synapses in CNS
are usually surrounded by the processes of astrocytes. Thus, astrocytes electrically insulate

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synapses and separate them from one another. However, the processes of astrocyte terminate on
blood vessel and axon.

b. Oligodendrocytes
Oligodendrocytes are found close to the myelinated axons in the brain and spinal cord.
Oligodendrocytes provide myelin sheath for neurons in CNS in a much similar way the Schwann
cells do for peripheral nerves. The processes of oligodendrocytes wrap many times around an
axon to form the myelin sheath. This sheath not only insulates axons from one another, but also
limits current flow across the axon membrane (axolemma). Because of this myelination, action
potential is conducted in a saltatory fashion in myelinated fibers, which is much faster than the
transmission of impulse in unmyelinated fibers.

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c. Microglia
Microglia are the smallest cells in the central nervous system. They are the scavenger
cells in brain. Thus, microglia contains phagocytic vesicle. If the nervous tissue is damaged or
infected, these cells enlarge and become mononuclear phagocytes to eliminate debris and
organisms.

d. Ependymal Cells
Ependymal cells line the surfaces of the brain’s ventricles and central canal of the
spinal cord. Their function is unclear.

Functions of Glial Cells


1. They form the mechanical matrix in which neurons are embedded.
2. They play metabolic and nutritive roles for neurons.
3. They may help to regulate blood flow through the brain.
4. They may act as a sink or source of ions.
5. They insulate axons and synapses and electrically isolate them from one another.
6. They phagocytose neural debris.

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RECEPTORS
Information about the internal and external environment activates the CNS via a variety
of sensory receptors. Receptors are sensory (afferent) nerve endings that terminate in periphery
as unmyelinated endings or specialized capsulated structures. Receptors give response to the
stimulus. When stimulated, receptors produce a series of impulses, which are transmitted through
the afferent nerves. Sensory receptors detect the state of the body or the state of the surroundings.
In nervous system, receptors act as a biological transducer. A transducer is a device that converts
one form of energy to another. Hence, receptors are often defined as the biological transducers,
which convert (transducer) various forms of energy (stimuli) in the environment into action
potentials in nerve fiber.

Classification of Receptors
Receptors are classified into two basic types, namely; Exteroceptors and Interoceptors.

1. Exteroceptors
Exteroceptors are the receptors, which give response to stimuli arising from outside the
body.

a. Cutaneous Receptors

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These are receptors situated in the skin. Cutaneous receptors are also called
mechanoreceptors because of their response to mechanical stimuli such as touch, pressure and
pain. Touch and pressure receptors give response to vibration also. Potentially harmful stimuli
such as extreme heat and extreme cold are said to be mediated by nociceptors.

b. Chemoreceptors
Receptors, which give response to chemical stimuli or changes in the chemical
composition of the environment is called the chemoreceptors.

c. Telereceptors
Telereceptors are the receptors that give response to stimuli arising away from the body.
These receptors are also called the distance receptors.

2. Interoceptors
Interoceptors are the receptors, which give response to stimuli arising from within the
body.

a. Proprioceptors
Proprioceptors are the receptors, which give response to change in the position of
different parts of the body in space at any given instant. However, the conscious component of
proprioception (“body image”) is actually synthesized from information coming not only from
receptors in and around joints but also from cutaneous touch and pressure receptors.

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b. Visceroceptors
Receptors situated in the viscera are called visceroceptors. This receptor senses such as
those sensitive to changes in the plasma level of O2, pH, and osmolality.

Properties of Receptors
1. Sensory Adaptation
Adaptation is the decline in discharge of sensory impulses when a receptor is stimulated
continuously with constant strength. It is also called sensory adaptation or desensitization.
Depending upon adaptation time, receptors are divided into two types:
i. Tonic receptors, which adapt slowly. Muscle spindle, pain receptors and cold
receptors are the tonic receptors.
ii. Phasic receptors, which get adapted rapidly. Touch and pressure receptors are the
phasic receptors.
2. Specificity of Response
Specificity of response or Müller law refers to the response given by a particular type of
receptor to a specific sensation. For example, pain receptors give response only to pain sensation.
Similarly, temperature receptors give response only to temperature sensation. In addition, each
type of sensation depends upon the part of the brain in which its fibers terminate.
3. Response to Increase in Strength of Stimulus
During the stimulation of a receptor, if the response given by the receptor is to be
doubled, the strength of stimulus must be increased 100 times. This phenomenon is called
Weber-Fechner law (R = k log S; where R = Intensity of response [sensation]; k = Constant S =
Intensity of stimulus), which states that intensity of response (sensation) of a receptor is directly
proportional to logarithmic increase in the intensity of stimulus.
4. Sensory Transduction
For receptor, sensory transduction refers to the process whereby the energy (stimulus) in
the environment is converted into action potentials (electrical impulses) in nerve fiber. Take for
example, when a receptor is stimulated, it responds by sending sensory information about the
stimulus to CNS via series of processes such as; the development of receptor potential in the
receptor cell and development of action potential in the sensory nerve. It is important to note that

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sensory transduction varies depending upon the type of receptor. For instance; touch receptor
converts mechanical energy into action potential in the sensory nerve fiber; while chemoreceptor
converts chemical energy into action potential in the sensory nerve fiber.
5. Receptor Potential
Receptor potential is a non-propagated transmembrane potential difference that
develops when a receptor is stimulated. It is also called generator potential that does not obey
all-or-none law. Receptor potential is not action potential. It is a graded potential that is short
lived and hence called transient receptor potential. It is similar to excitatory postsynaptic
potential (EPSP) in synapse, endplate potential in neuromuscular junction and electrotonic
potential in the nerve fiber.
6. Law of Projection
When a sensory pathway from receptor to cerebral cortex is stimulated on any particular
site along its course, the sensation caused by stimulus is always felt (referred) at the location of
receptor, irrespective of site stimulated. This phenomenon is known as law of projection.
Examples of law of projection; If somesthetic area in right cerebral cortex, which receives
sensation from left hand is stimulated, sensations are felt in left hand and not in head. Another
example is the sensation complained by amputated patients in the missing limb with scar
formation at the cut end (phantom limb) is the best example of law of projection. The patient
feels as if the sensation is originating from nonexistent leg on if the cut end of sensory fibers are
stimulated during movement of thigh.

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SYNAPSE
The junction of an axonal ending with a nerve cell, a muscle cell, or a glandular cell is called a
synapse. It is the junction between two neurons. It is not an anatomical but physiological
continuity between two nerve cells. At synapses the propagated action potential is transmitted to
the next cell.

Classification of Synapses
1. Anatomical classification
Synapse can be classified into three types based on their anatomy:
a. Axoaxonic synapse: this synapse is formed when axon of one neuron terminates on axon
of another neuron

b. Axodendritic synapse: this synapse is formed when the axon of one neuron terminates
on dendrite of another neuron

c. Axosomatic synapse: this synapse is formed when axon of one neuron ends on soma
(cell body) of another neuron

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2. Physiological classification
Synapse is classified into two types based on their functions. However, generally the word
synapse refers to a chemical synapse;

a. Electrical Synapse: Axon always forms a "gap junction” that is in closest contact with the
nerve cell on which it ended so that the propagated impulse could be transmitted without
interruption to another nerve cell. Thus, electrical synapse forms a physiological continuity
between the presynaptic and the post-synaptic neurons provided by gap junction between
the two neurons. There is direct exchange of ions between the two neurons through the gap
junction, hence, synaptic delay is very less because of the direct flow of current and the
impulse is transmitted in either direction. This type of impulse transmission occurs in some
tissues like; the cardiac muscle fibers, smooth muscle fibers of intestine and the epithelial
cells of lens in the eye.

b. Chemical Synapse: The axonal ending when stimulated releases a chemical substance that
produces an excitatory or an inhibitory effect at the neighboring cell membrane. This type of
synapse is called a chemical synapse. There are two neurons taking part at a synapse. These
two neurons are known as pre-synaptic neuron and post synaptic neuron. These two
membranes are known as pre-synaptic membrane and post-synaptic membrane. In this type
of functional synapse, there is no continuity between the two neurons because of the presence
of a space called synaptic cleft. The gap is about 20–50 nm (nano meter). Inside the pre-
synaptic membrane there are several vesicles (sacs) that are filled with neurotransmitters.
There are three kinds of synaptic vesicles: small, clear synaptic vesicles that contain
acetylcholine, glycine, GABA, or glutamate; small vesicles with a dense core that contain
catecholamines; and large vesicles with a dense core that contain neuropeptides. An action
potential or an impulse reaches the tip of pre synaptic neuron. This makes the vesicles to

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move towards pre-synaptic membrane. Action potential reaching the presynaptic terminal
causes release of neurotransmitter substance from the vesicles of this terminal. In the
presence of calcium ions the vesicle ruptures and releases the neurotransmitter into the
synaptic cleft. The neurotransmitter travels across the synaptic cleft and reach the post
synaptic membrane. Here it binds with receptor protein. This binding changes the
permeability of post synaptic membrane. The membrane becomes more permeable to sodium
ions. Na+ influx takes place. Post synaptic membrane gets depolarised. This leads to the
generation of fresh action potential in the post synaptic neuron. The neurotransmitters
released at the synaptic cleft reach the post synaptic membrane and produces the desired
change in the post synaptic potential. After that the neurotransmitters are inactivated by the
following means:
i. Most of the neurotransmitters are actively taken back by the pre-synaptic membrane
and are then reused again and again. This is called transmitter re-uptake. It occurs
especially prominently at the presynaptic terminals of the sympathetic nervous
system. Reuptake process involves a specific carrier protein for each
neurotransmitter; for example; the re-uptake of norepinerphrine.
ii. The neurotransmitters are degraded enzymatically within the cleft itself. For instance,
in the case of acetylcholine, the enzyme cholinesterase is present in the cleft, bound in
the proteoglycan matrix that fills the space. Similar effects occur for other
transmitters.
iii. Small quantity of neurotransmitter escapes at the interstitial space (Surrounding
fluids) by diffusion.

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Fig: Structure of chemical synapse

Functions of Synapse
Main function of the synapse is to transmit the impulses, i.e. action potential from one
neuron to another. Some of the synapses inhibit these impulses. So the impulses are not
transmitted to the postsynaptic neuron. Hence, the functions of synapses are divided into two
types: Excitatory synapses, which transmit the impulses (excitatory function) and Inhibitory
synapses, which inhibit the transmission of impulses (inhibitory function).
A. EXCITATORY FUNCTION
Excitatory Postsynaptic Potential
Excitatory postsynaptic potential (EPSP) is the non-propagated electrical potential that
develops during the process of synaptic transmission. When the action potential reaches the pre-
synaptic axon terminal, the voltage gated calcium channels at the pre-synaptic membrane are
opened. Now, the calcium ions enter the axon terminal from ECF causing the release of
neurotransmitter substance from the vesicles by means of exocytosis. Neurotransmitter (NT),
which is excitatory in function (excitatory neurotransmitter) passes through pre-synaptic
membrane and synaptic cleft and reaches the postsynaptic membrane. The NT binds with
receptor protein present in postsynaptic membrane to form neurotransmitter receptor complex.
NT receptor complex causes production of a non-propagated EPSP triggers the opening of

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voltage-gated sodium channels and influx of sodium ions with development of mild
depolarization.

B. INHIBITORY FUNCTION
Synaptic inhibition in CNS limits the number of impulses going to muscles and enables
the muscles to act properly and appropriately. Thus, the inhibition helps to select exact number
of impulses and to omit or block the excess ones. Inhibition of synaptic transmission is classified
into five types:

1. Presynaptic or Indirect Inhibition


Pre-synaptic inhibition occurs due to the failure of pre-synaptic axon terminal to release
sufficient quantity of excitatory neurotransmitter substance. It is also called indirect inhibition.
Presynaptic inhibition is mediated by axoaxonal synapses. It is prominent in spinal cord and
regulates the propagation of information to higher centers in brain. Normally, during synaptic
transmission, action potential reaching the presynaptic neuron produces development of EPSP in
the postsynaptic neuron. But, in spinal cord, a modulatory neuron called presynaptic
inhibitory neuron forms an axoaxonic synapse with the presynaptic neuron.

2. Postsynaptic or Direct Inhibition


Postsynaptic inhibition is the type of synaptic inhibition that occurs due to the release of
an inhibitory neurotransmitter from pre-synaptic terminal instead of an excitatory
neurotransmitter substance. It is also called direct inhibition. Inhibitory neurotransmitters are
gamma aminobutyric acid (GABA), dopamine and glycine. Inhibitory postsynaptic potential
(IPSP) is the electrical potential in the form of hyperpolarization that develops during
postsynaptic inhibition. Inhibitory neurotransmitter substance acts on postsynaptic membrane by
binding with receptor. Transmitter receptor complex opens the ligand-gated potassium channels
instead of sodium channel. The potassium ions, which are available in plenty in the cell body of
postsynaptic neuron move to ECF. Simultaneously, chloride channels also open and chloride
ions (which are more in ECF) move inside the cell body of postsynaptic neuron leading to
hyperpolarization.

3. Renshaw Cell or Negative Feedback Inhibition

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Negative feedback inhibition is the type of synaptic inhibition, which is caused by


Renshaw cells in spinal cord. Renshaw cells are small motor neurons present in anterior gray
horn of spinal cord. Anterior nerve root consists of nerve fibers, which leave the spinal cord.
These nerve fibers arise from αmotor neurons in anterior gray horn of the spinal cord and reach
the effector organ, muscles. Some of the fibers called collaterals fibers terminate on Renshaw
cells instead of leaving the spinal cord. Stimulation of the Renshaw cell sends inhibitory
impulses to αmotor neurons so that, the discharge from motor neurons is reduced.

4. Feedforward Inhibition
Feedforward synaptic inhibition occurs in cerebellum and it controls the neuronal
activity in cerebellum. During the process of neuronal activity in cerebellum, stellate cells and
basket cells, which are activated by granule cells, inhibit the Purkinje cells by releasing GABA.
This type of inhibition is called feedforward inhibition.

5. Reciprocal Inhibition
Inhibition of antagonistic muscles when a group of muscles are activated is called
reciprocal inhibition. It is because of reciprocal innervation.

PROPERTIES OF SYNAPSE
1. Electrical properties: Synapse has both EPSP and IPSP properties.
2. Uni directionality: Impulses are transmitted only in one direction in synapse (Bell-
Magendie law), i.e. from pre-synaptic neuron to post-synaptic neuron.
3. Synaptic Delay: it is a short delay that occurs during the transmission of impulses
through the synapse. It is due to the time taken (0.3 to 0.5 millisecond) for
neurotransmitter to be released, neurotransmitter to be transported from axon terminal to
postsynaptic membrane and action of the neurotransmitter to open the ionic channels in
postsynaptic membrane.
4. Summation: it involves the fusion of effects or progressive increase in the excitatory
postsynaptic potential in post synaptic neuron when many pre-synaptic excitatory
terminals are stimulated simultaneously or when single pre-synaptic terminal is
stimulated repeatedly. Increased EPSP triggers the axon potential in the initial segment of

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axon of postsynaptic neuron. It can either be spatial summation (simultaneous stimulation


of many pre-synaptic terminals) or Temporal summation (repeated stimulation of one
pre-synaptic terminals).
5. Fatigue: Fatigue at synapse is due to the depletion of neurotransmitter substance,
acetylcholine that are destroyed by acetylcholinesterase. During continuous muscular
activity, synapse becomes the seat of fatigue along with Betz cells present in motor area
of frontal lobe of cerebral cortex.

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NEUROTRANSMITTERS
Neurotransmitters (NT) are the chemical substances liberated at the nerve endings and help to
transfer the message of the nerve impulses in the presynaptic neuron to an adjacent cell. NT acts
as a mediator for the transmission of nerve impulse from one neuron to another neuron through
a synapse. e.g., postsynaptic neuron, a muscle or a gland. Otto Loewi an Austrian scientist in
1921 was the first to discover the existence of NT while he was working on the Heart of frog.
Loewi practically experimented the stimulation of the vagus nerve of frogs and came out with
this discovery of a NT called Ach.

Properties of NT
1. The chemical must be synthesised in the neuron concerned.
2. Should be stored in the presynaptic endings.
3. Should be released at the synapse.
4. Should have its specific receptors on the post synaptic membrane.
5. On binding with receptor proteins on the surface of the post synaptic membrane, it should
produce either EPSP or IPSP.
6. Should be disposed quickly by a suitable system as soon as its action is over.
7. Exogenous application of the substance should mimic the effects of stimulation of the neuron.

Types of Neurotransmitters
There are about 40 different neurotransmitters. Identified neurotransmitters and neuromodulators
can be divided into two major categories: small-molecule transmitters and large-molecule
transmitters;
1. Small molecule, rapidly acting transmitters- which cause acute response:
a. Aminoacids (Neurotransmitters of this group are inhibitory and excitatory in action) e.g:
(i) Gamma-amino butyric acid (GABA)
(ii) Glycine
(iii) Glutamate (glutamic acid)
(iv) Aspartate (aspartic acid)
b. Different amines (Neurotransmitters of this group are the modified amino acids i.e
monoamine) e.g:

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(i) Norepinephrine
(ii) Epinerphrine
(iii) Dopamine
(iv) Serotonin
(v) Histamine

2. Large-molecule, slowly acting transmitters having prolonged affect:


a. Neuropeptides: Usually neuropeptides are colocalized with one of the small-molecule
neurotransmitters.
(i) neuroactive peptides- releasing hormones from hypothalamus: TRH, LH releasing hormone,
somotostatin.
(ii) Pituitary peptides: ACTH, beta endorphins, vasopressin, oxytocin.
(iii) Peptides acting on gut and brain: leucin, enkephalin, methionine, substance P,
cholecystokinin, Vasoactive intestinal polypeptide (VIP), neurotensin, insulin, glucagon.
(iv) Neuropeptides from other tissues: angiotensin II, bradykinin, carnosine, bombesin.

3. Other Substances
Some neurotransmitters do not fit into any of these categories. One such substance is
acetylcholine. It is formed from the choline and acetyl coenzyme A in the presence of the
enzyme called choline acetyltransferase. There are also other substances thought to be released
into the synaptic cleft to act as either a transmitter or modulator of synaptic transmission. These
include purine derivatives like adenosine and adenosine triphosphate (ATP) and a gaseous
molecule, nitric oxide (NO).

Excitatory and inhibitory neurotransmitters


Excitatory Inhibitory Neurotransmitters with excitatory
neurotransmitters
Neurotransmitters and inhibitory actions
1. Acetylcholine 1. Gamma-aminobutyric acid 1. Noradrenaline
2. Nitric oxide 2. Glycine 2. Adrenaline
3. Histamine 3. Dopamine
4. Glutamate 4. Serotonin
5. Aspartate

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ACETYLCHOLINE (Excitatory)
Acetylcholine is a cholinergic neurotransmitter. It possesses excitatory function. It
produces the excitatory function by opening the ligand-gated sodium channels. Ach is
synthesized in the cholinergic nerve endings. Synthesis takes place in axoplasm and Ach is
stored in the vesicles. It is synthesized from acetyl coenzyme A (acetyl CoA). It combines with
choline in the presence of the enzyme choline acetyltransferase to form Ach. Acetylcholine is the
transmitter substance at the neuromuscular junction and synapse. It is also released by the
following nerve endings: Preganglionic sympathetic nerve, Postganglionic sympathetic
cholinergic nerves, Preganglionic parasympathetic nerve, Postganglionic parasympathetic nerve,
Nerves in amacrine cells of retina and Many regions of brain. Action of Ach is short lived.
Within one millisecond after the release from the vesicles, it is hydrolyzed into acetate and
choline by the enzyme acetylcholinesterase (This enzyme is present in basal lamina of the
synaptic cleft). There are two types of receptors through which Ach acts on the tissues namely,
muscarinic receptors and nicotinic receptors.

SEROTONIN (Inhibitory)
Serotonin is also known as 5-hydroxytryptamine (5-HT). Serotonin is formed in the
body by hydroxylation and decarboxylation of the essential amino acid tryptophan. After release
from serotonergic neurons, much of the released serotonin is recaptured by an active reuptake
mechanism and inactivated by monoamine oxidase (MAO) to form 5-hydroxyindoleacetic acid
(5-HIAA). Large amount of serotonin (90%) is found in enterochromatin cells of GI tract. Small
amount is found in platelets and nervous system. It is secreted in the following structures:
hypothalamus, limbic system, cerebellum, spinal cord, dorsal raphe nucleus of midbrain, GI
tract, retina, lungs and platelets. The number of cloned and characterized serotonin receptors has
increased rapidly. There are at least seven types of 5-HT receptors (from 5-HT1 through 5-HT7
receptors). Some of the serotonin receptors are presynaptic, and others are postsynaptic.
Serotonin is an inhibitory substance. It inhibits impulses of pain sensation in posterior gray horn
of spinal cord. It is supposed to cause depression of mood and sleep. Serotonin causes
vasoconstriction, plate let aggregation and smooth muscle contraction. It also controls food
intake.

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NEUROMODULATORS
Neuromodulator is the chemical messenger, which modifies and regulates activities that
take place during the synaptic transmission. Generally the neuromodulators are peptides. So
neuro-modulators are often referred as neuropeptides. Almost all the peptides found in nervous
tissues are neuromodulators. These peptides do not propagate nerve impulses like
neurotransmitters. Neuromodulators are classified into two types namely; Opioid peptides and
Non-opioid peptides.

a. Opioid Peptides
Peptides, which bind to opioid receptors are called opioid peptides. Opioid peptides are also
called opioid neuropeptides or opioid neuromodulators. Opioid receptors are of three types μ, к
and δ. These proteins are called opioid receptors because of their affinity towards the opiate or
morphine, which are derived from opium. Opioid peptides are of three types:
i. Enkephalins- are secreted in many parts of brain, substantia gelatinosa and retina
(Inhibit pain sensation)
i. Dynorphins: are secreted in the hypothalamus, posterior pituitary and duodenum
(inhibit pain sensation)
ii. Endorphins: are secreted in the Thalamus, hypothalamus, brainstem and retina
(inhibit pain sensation)

b. Non-opioid Peptides
Non-opioid neuropeptides act by binding with G-protein coupled receptors. These
neuropeptides are also called non-opioid neuromodulators. Non-opioid peptides are;
i. Bradykinin – secreted in Blood vessels, kidneys (Causes vasodilatation)
ii. Secretin - secreted in Cerebral cortex, hypothalamus, thalamus, olfactory bulb,
brainstem and small intestine; they inhibits gastric secretion and motility)
iii. Substance P –secreted in Brain, spinal cord, retina peripheral nerves and intestine
(Mediates pain, Causes vasodilatation, regulates anxiety, stress, mood disorders,
neurotoxicity, nausea and vomiting.

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iv. Cholecystokinin (CCK) – secreted in hypothalamus, cerebral cortex, retina and


small intestine (Contracts gallbladder, Increases intestinal motility and Inhibits gastric
motility)
v. Gastrin: secreted in GI tract, Hypothalamus, medulla oblongata, posterior pituitary
and (Stimulates islets in pancreas, Increases gastric secretion and motility)
vi. Motilin: secreted Cerebral cortex, cerebellum, posterior pituitary and intestine
(Stimulates intestinal motility)
vii. Neuropeptide Y: secreted in the Medulla, hypothalamus and small intestine
(Increases food intake, causes vasoconstriction and increases enteric blood flow)
viii. Vasoactive intestinal polypeptide: secreted in Cerebral cortex, hypothalamus, retina
and intestine (Causes vasodilatation)
ix. Vasopressin: secreted in posterior pituitary, medulla oblongata and spinal cord
(Causes vasoconstriction)
x. Oxytocin: secreted in posterior pituitary, medulla oblongata and spinal cord
(Stimulates milk ejection and uterine contraction)
xi. Corticotropin-releasing hormone (CRH): secreted in Hypothalamus (Stimulates
release of ACTH)
xii. Growth hormone-releasing polypeptide: secreted in Hypothalamus (Stimulates
release of GHRH)
xiii. Thyrotropin-releasing hormone (TRH): secreted in Hypothalamus (Stimulates
release of thyroid hormones)
xiv. Gonadotropin-releasing hormone (GnRH): secreted in Hypothalamus (Stimulates
release of growth hormone)
xv. Somatostatin: secreted in Hypothalamus, other parts of brain, substantia gelatinosa
and retina (Inhibits growth hormone secretion Decreases food intake)
xvi. Angiotensin II: secreted in the hypothalamus, brainstem and spinal cord (Causes
vasoconstriction)
xvii. Endothelin: secreted in Posterior pituitary, brainstem and endothelium (Causes
vasoconstriction)

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xviii. Ghrelin: secreted in Hypothalamus, stomach, pituitary, kidney and placenta


(Promotes GH release Induces appetite and food intake and stimulates gastric
emptying)

Differences between Neurotransmitters and Neuromodulators


Neurotransmitters Neuromodulators
Propagate nerve impulse through synapse Modify and regulate synaptic transmission
Packed in small synaptic vesicles Packed in large synaptic vesicles
Generally, neuron has only one Neuron may have one or more
neurotransmitter neuromodulators
Found only in axon terminals Found in all parts of neuron like soma,
dendrite, axon and nerve endings.
Act by changing the electric potential – Have diverse actions
depolarization or repolarization
Chemically, neurotransmitters are amino Chemically, neuromodulators are only
acids, amine or others peptides

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REFLEX ACTIVITY AND REFLEX ARC


Reflex action is the involuntary response to a stimulus. That is, reflex activity is the
response to a peripheral nervous stimulation that occurs without our consciousness. It is a type of
protective mechanism and it protects the body from irreparable damages; E.g touching a hot
object or when a bright light comes in contact with the eye. The basic unit of integrated reflex
activity is the reflex arc. That is, the anatomical nervous pathway for a reflex action. A simple
reflex arc includes five components namely; a sense organ or receptor that receives the stimulus,
an afferent or sensory neuron that transmits sensory impulses from the receptor to the center
which is located in the brain or spinal cord (The center receives the sensory impulses via afferent
nerve fibers and in turn, it generates appropriate motor impulses), an efferent or motor neuron
that transmits motor impulses from the center, and an effector organ which is the structure such
as muscle or gland where the activity occurs in response to stimulus.

Figure: Simple reflex arc

In mammals, the connection between afferent and efferent somatic neurons is generally
in the brain or spinal cord. The afferent neurons enter via the dorsal roots or cranial nerves and
have their cell bodies in the dorsal root ganglia or in the homologous ganglia on the cranial
nerves. The efferent fibers leave via the ventral roots or corresponding motor cranial nerves. The
principle that in the spinal cord the dorsal roots are sensory and the ventral roots are motor is
known as the Bell–Magendie law.
Activity in the reflex arc starts in a sensory receptor with a receptor potential whose
magnitude is proportional to the strength of the stimulus. This generates all-or-none action

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potentials in the afferent nerve, the number of action potentials being proportional to the size of
the generator potential. In the central nervous system (CNS), the responses are again graded in
terms of excitatory postsynaptic potentials (EPSPs) and inhibitory postsynaptic potentials
(IPSPs) at the synaptic junctions. All-or-none responses are generated in the efferent nerve.
When these reach the effector, they again set up a graded response. When the effector is smooth
muscle, responses summate to produce action potentials in the smooth muscle, but when the
effector is skeletal muscle, the graded response is always adequate to produce action potentials
that bring about muscle contraction.

Classification of Reflexes
Reflexes are classified by six different methods based on the following factors;
a. Depending Upon Whether Inborn or Acquired Reflexes
i. Unconditioned or Inborn Reflexes
This class of reflex is known as natural reflexes or unconditioned reflex, i.e they are
present from since the time of birth. Such reflexes do not require previous learning, training or
conditioning. Best example is the secretion of saliva when a drop of honey is kept in the mouth
of a newborn baby for the first time. The baby does not know the taste of honey, but still saliva is
secreted.

ii. Conditioned Reflexes or Acquired Reflexes


Acquired or conditioned reflexes are the reflexes that are developed after conditioning
or training. These reflexes are not inborn but, acquired after birth. Such reflexes need previous
learning, training or conditioning. Example is the secretion of saliva by sight, smell, thought or
hearing of a known edible substance.

b. Depending Upon Situation – Anatomical Classification


Reflexes can be classified depending upon the situation of the center.
i. Cortical Reflexes: these reflexes are the reflexes that have their center in cerebral cortex
ii. Cerebellar Reflexes: these reflexes are the reflexes which have their center in cerebellum.
iii. Midbrain Reflexes: these reflexes are the reflexes which have their center in midbrain.
iv. Medullary or Bulbar Reflexes: these reflexes are the reflexes which have their center in
medulla oblongata.

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v. Spinal Reflexes: Reflexes having their center in the spinal cord are called spinal reflexes.
Depending upon the segments involved, spinal reflexes are divided into three groups:
 Segmental spinal reflexes
 Intrasegmental spinal reflexes
 Suprasegmental spinal reflexes.

c. Depending Upon the Number of Synapse


Depending upon the number of synapse in reflex arc, reflexes are classified into two
types:
i. Monosynaptic Reflexes
Reflexes having only one synapse in the reflex arc are called monosynaptic reflexes.
Stretch reflex is the best example for monosynaptic reflex and it is elicited due to the stimulation
of muscle spindle.

ii. Polysynaptic Reflexes


Reflexes having more than one synapse in the reflex arc are called polysynaptic reflexes.
Flexor or withdrawal reflexes are the polysynaptic reflexes.

d. Depending Upon Purpose:


This method reflexes is known as physiological classification which is depending upon
the purpose (functional significance);

i. Flexor or Protective Reflexes


These are the reflexes which protect the body from nociceptic (harmful) stimuli. These
reflexes are also called withdrawal reflexes or flexor reflexes. Protective reflexes involve
flexion at different joints hence the name flexor reflexes

ii. Extensor or Antigravity Reflexes


These are the reflexes that protect the body against gravitational force. These reflexes
are also called the extensor reflexes because, the extensor muscles contract during these reflexes
resulting in extension at joints.

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e. Depending Upon Whether Somatic or Visceral Reflexes


i. Somatic Reflexes
Somatic reflexes are the reflexes, for which the reflex arc is formed by somatic nerve
fibers. These reflexes involve the participation of skeletal muscles. And there may be flexion or
extension at different joints during these reflexes.

ii. Visceral or Autonomic Reflexes


Visceral or autonomic reflexes are the reflexes, for which at least a part of reflex arc is
formed by autonomic nerve fibers. These reflexes involve participation of smooth muscle or
cardiac muscle. Visceral reflexes include pupillary reflexes, gastrointestinal reflexes,
cardiovascular reflexes, respiratory reflexes, etc. It is important to note that some reflexes like;
swallowing, coughing or vomiting are considered as visceral reflexes. However, these reflexes
involve some participation of skeletal muscles.

f. Depending Upon Clinical Basis


Depending upon the clinical basis, reflexes are classified into four types:
i. Superficial Reflexes: these are the reflexes, which are elicited from the surface of the
body. Superficial reflexes are of two types: mucus membrane reflexes and cutaneous
or skin reflexes.
ii. Deep Reflexes: they are elicited from deeper structures beneath the skin like tendon.
These reflexes are otherwise known as tendon reflexes.
iii. Visceral Reflexes: they are the reflexes arising from pupil and visceral organs.
Examples of visceral reflexes:
 Pupillary reflexes: are the reflexes in which, the size of pupil is altered.
Light reflex: Direct light reflex and Indirect or consensual light reflex
Accommodation reflex: Constriction of pupil, Increase in anterior curvature
of lens and Convergence of eyeball
Ciliospinal reflex: the dilatation of pupil due to stimulation of skin over the
neck.
 Oculocardiac reflexes: are the reflexes in which, heart rate decreases due to
the pressure applied over eyeball.

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 Carotid sinus reflex: are the reflexes in which heart rate and blood pressure
decreases due to pressure over carotid sinus in neck due to tight collar.
iv. Pathological Reflexes; they are reflexes that are elicited only in pathological
conditions. Well-known pathological reflexes are:
 Babinski sign: this is abnormal plantar reflex named after the discoverer
Joseph Babinski. In this reflex, a gentle scratch over the outer edge of the
sole of foot causes plantar flexion and adduction of all toes.
 Clonus: this is a series of rapid and repeated involuntary jerky movements,
which occur while eliciting a deep reflex. Examples; Ankle Clonus and
Patellar Clonus
 Pendular movement: are the slow oscillatory movements (instead of brisk
movements) that are developed while eliciting a tendon jerk. Unlike clonus,
pendular movements occur because of hypotonicity of muscles.

Properties of Reflexes
i. One Way Conduction (Bell-Magendie Law): During any reflex activity, impulses
are transmitted in only one direction through the reflex arc as per Bell- Magendie
law. The impulses pass from receptors to center and then from center to effector
organ.
ii. Summation: in this regards, reflexes can either exhibit spatial summation or temporal
Summation. Spatial Summation involves when two afferent nerve fibers supplying a
muscle are stimulated separately with subliminal stimulus, there is no response. But
the muscle contracts when both the nerve fibers are stimulated together with same
strength of stimulus. Temporal Summation involves when one nerve fiber is
stimulated repeatedly with subliminal stimuli, these stimuli are summed up to give
response in the muscle.
iii. Reaction Time: this is the time interval between application of stimulus and the
onset of reflex. It depends upon the length of afferent and efferent nerve fibers,
velocity of impulse through these fibers and central delay. Central delay is the delay
at the synapse. It is also called synaptic delay.

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iv. Recruitment: this is defined as the successive activation of additional motor units
with progressive increase in force of muscular contraction. Recruitment is similar to
the effect of temporal summation.
v. Occlusion: this is demonstrated in a flexor reflex involving a muscle, which is
innervated by two motor nerves. When both nerves are stimulated simultaneously, the
tension developed by the muscle is less than the sum of the tension developed when
each nerve is stimulated separately. Occlusion is due to the overlapping of the nerve
fibers during the distribution.

vi. Subliminal Fringe: In some reflexes involving the muscle with two nerve fibers, the
tension developed by simultaneous stimulation of two nerves is greater than the sum
of tension produced by the stimulation of these nerves separately. It is due to the
effect of spatial summation.

vii. Fatigue: When a reflex activity is continuously elicited for a long time, the response
is reduced slowly and at one stage, the response does not occur. This type of failure to
give response to the stimulus is called fatigue. Center or the synapse of the reflex arc
is the first seat of fatigue.
viii. Rebound Phenomenon: Reflex activities can be forcefully inhibited for some time.
But, when the inhibition is suddenly removed, the reflex activity becomes more
forceful than before inhibition.

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SPINAL CORD
Anatomy of the Spinal Cord

Figure 1: Section of spinal cord: thoracic segment

 Spinal cord is a long fragile tube-like band or cylindrical shaped tissue, nerve and cell
that lies loosely through the center of the spine (the vertebral canal). The length of the
spinal cord is about 45 cm in males and about 43 cm in females. It is 2cm wide.
 It extends from the base of the skull (foramen magnum) where it is continuous with
medulla oblongata, above and up to the lower border of first lumbar vertebra below.
 The vertebral column covers and protects the spinal cord. The bones in the vertebral
column are called vertebrae (the stalk on top of each other from the pelvic bone to the
skull).
 Spinal cord has two spindle-shaped swellings, namely cervical and lumbar
enlargements. These two portions of spinal cord innervate upper and lower extremities
respectively.
 Below the lumbar enlargement, spinal cord rapidly narrows to a cone-shaped termination
called conus medullaris. A slender non-nervous filament called filum terminale extends

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from conus medullaris downward to the fundus of the dural sac at the level of second
sacral vertebra.
 Three layers or sheaths called meninges which are membranous in nature surrounds the
spinal cord, namely; Dura mater (has 2 layers; Periosteal and meningeal with epidural
space between them), Pia mater and Arachnoid mater (Subarchnoid space lies
between). These coverings continue as coverings of brain. Meninges are responsible for
protection and nourishment of the nervous tissues.
 The space at the end of spinal cord, especially between L1 and L2 vertebral segments is
used in tapping cerebrospinal fluid during lumbar puncture.
 Spinal cord is divided into 5 parts which makes up 31 segments each having a 31
bilaterally symmetrical pairs of spinal nerves (motor and a sensory nerve root),
namely;
1. Cervical segments/Cervical spinal nerves = 8
2. Thoracic segments/Thoracic spinal nerves = 12
3. Lumbar segments/Lumbar spinal nerves = 5
4. Sacral segments/Sacral spinal nerves = 5
5. Coccygeal segment/Coccygeal spinal nerves = 1

 Each spinal nerve arises from different regions of the vertebral column and are named
accordingly (Neck, Chest, Pelvic and Abdomen). It is formed by an anterior (ventral
motor) root and a posterior (dorsal sensory) root. Both the roots on either side leave
the spinal cord and pass through the corresponding intervertebral foramina.
 On the anterior surface of spinal cord, there is a deep furrow known as anterior median
fissure. Depth of this fissure is about 3 mm. Lateral to the anterior median fissure on
either side, there is a slight depression called the anterolateral sulcus. It denotes the exit
of anterior nerve root. On the posterior aspect, there is a depression called posterior
median sulcus. This sulcus is continuous with a thin glial partition called the posterior
median septum. It extends inside the spinal cord for about 5 mm and reaches the gray
matter.
 On either side, lateral to posterior median sulcus, there is posterior intermediate sulcus.
It is continuous with posterior intermediate septum, which extends for about 3 mm into

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the spinal cord. Lateral to the posterior intermediate sulcus, is the posterolateral sulcus.
This denotes the entry of posterior nerve root.

Function of the Spinal Cord


 The main function of spinal cord is to receive sensory inputs from peripheral structures
via somatic nerves and transmit them to the brain, and convey the signals originating
from brain motor and autonomic areas to the target structures (i.e it forms a vital link
between the brain and the body aiding primary processing of information).
 The spinal cord provides structural support and builds the body posture by connecting the
brain to the lower back. It is a delicate structure that contains nerve bundles and cells that
carries messages or nerve signals from the brain to the different part of the body vis
versa. This nerve signals aids sensations and facilitate flexible body movement. It reports
senses to the senses to the brain and manages reflexes.
 Understanding the physiology of the spinal cord helps in detecting and determining the
various methods to deal with diseases and dame related to the spinal cord. Thus, any
damage to the spinal cord affects movement and bodily functions.

GRAY MATTER OF SPINAL CORD


Gray matter of spinal cord is the collection of nerve cell bodies, dendrites and parts of
axons. It is placed centrally in the form of wings of the butterfly and it resembles the letter ‘H’.
Exactly in the center of gray matter, there is a canal called the spinal canal and is filled with
CSF. Ventral and the dorsal portions of each lateral half of gray matter are called ventral
(anterior) and dorsal (posterior) gray horns respectively. In addition, the gray matter forms a
small projection in between the anterior and posterior horns in all thoracic and first two lumbar
segments. It is called the lateral gray horn. Part of the gray matter anterior to central canal is
called the anterior gray commissure and part of gray matter posterior to the central canal is
called posterior gray commissure.

Neurons in Gray Matter of Spinal Cord

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1. Golgi type I neurons: Golgi type I neurons have long axons and are usually found in
anterior horns. Axons of these neurons form the long tracts of spinal cord.
2. Golgi type II neurons: Golgi type II neurons have short axons, which are found mostly
in posterior horns. Axons of these neurons pass towards the anterior horn of same side or
opposite side.

Organization of Neurons in Gray Matter


Organization of neurons in the gray matter of spinal cord is described in two methods: Nuclei
or columns and Laminae or layers.

Figure 2: Neurons in gray horn of spinal cord: thoracic segment

1. Nuclei
Clusters of neurons are present in the form of nuclei or cell columns in gray matter.
a. Nuclei in Posterior Gray Horn: Posterior gray horn contains the nuclei of sensory
neurons, which receive impulses from various receptors of the body through posterior
nerve root fibers. There are four types of nuclei of sensory neurons

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 Marginal nucleus is also called posteromarginal nucleus, marginal zone


nucleus or border nucleus
 Substantia gelatinosa of Rolando
 Chief sensory nucleus or nucleus proprius
 Dorsal nucleus of Clarke or Clarke column of cells
b. Nuclei in Lateral Gray Horn: Lateral gray horn has cluster of neurons called
intermediolateral nucleus. The neurons of this nucleus give rise to sympathetic
preganglionic fibers, which leave the spinal cord through the anterior nerve root. Inter m
ediolateral nucleus extends between T1 and L2 segments of spinal cord.
c. Nuclei in Anterior Gray Horn: Anterior gray horn contains the nuclei of lower motor
neurons, which are involved in motor function. These nuclei are present in almost all the
levels of spinal cord. Three types of motor neurons are present in lower motor neuron
nuclei:
 Alpha motor neurons (large and multipolar cells) ends in group of skeletal
muscles called extrafusal fibers.
 Gamma motor neurons (smaller cells scattered among alpha motor neurons) send
axons to intrafusal fibers of the muscle spindle.
 Renshaw cells (small cells) play an important role in synaptic inhibition

2. Laminae
Neurons of gray matter are distributed in laminae or layers. Each lamina consists of
neurons of different size and shape. This laminae was named by Brian Burke and Rexed in
1950 to be 10 in number.
a. Laminae in Posterior Gray Horn: Laminae I to VI constitute the posterior gray horn.
These laminae contain nuclei of sensory neurons, which are concerned with sensory
functions.
 Nuclei present in the laminae of posterior gray horn
 Marginal nucleus: Lamina I
 Substantial gelatinosa of Rolando: Laminae II and III
 Chief sensory nucleus : Laminae III, IV and V
 Dorsal nucleus of Clarke : Lamina VI

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b. Lamina in Lateral Gray Horn: Lateral gray horn contains only one lamina, the lamina
VII. It contains intermediolateral nucleus.
c. Laminae in Anterior Gray Horn: Laminae VIII and IX form the anterior gray horn.
These laminae contain nuclei of motor neurons, which are concerned with motor
functions.
d. Lamina Around Central Canal: Lamina X is found around the center of the spinal canal.
It contains neuroglia, which form the supporting tissue.

WHITE MATTER OF SPINAL CORD


It is formed by the bundles of both myelinated and nonmyelinated fibers, but
predominantly the myelinated fibers (i.e collections of axons permitting communication between
different layers of CNS). Anterior median fissure and posterior median septum divide the entire
mass of white matter into two lateral halves. The band of white matter lying in front of anterior
gray commissure is called anterior white commissure. Each half of the white matter is divided
by the fibers of anterior and posterior nerve roots into three white columns or funiculli.

I. Anterior or Ventral White Column or funiculus.


II. Lateral White Column or funiculus.
III. Posterior or Dorsal White Column or funiculus.

TRACTS IN SPINAL CORD


A tract is a collection of axons that carries specialized information. These groups of nerve fibers
passing through spinal cord are known as tracts of the spinal cord. The spinal tracts are divided
into two main groups. They are: Short and the long tracts.

1. Short Tracts: this involves fibers of the short tracts connect different parts of spinal cord
itself. They are of two types, namely:
a. Intrinsic or association tracts, which connect adjacent segments of spinal cord on the same
side
b. Commissural tracts, which connect opposite halves of same segment of spinal cord.

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2. Long Tracts: the long tracts or projection tracts of spinal cord connects the spinal cord with
other parts of central nervous system. They are of two types, namely:
a. Ascending tracts, which carry sensory impulses from the spinal cord to brain
b. Descending tracts, which carry motor impulses from brain to the spinal cord.

ASCENDING TRACTS OF SPINAL CORD


Ascending tracts of spinal cord carry the impulses of various sensations from the peripheral
structures to the different areas in the brain, mainly via thalamus to the sensory cortex. These
pathways are simply a set of neurons arranged in series.

Arrangement of Ascending Tracts


Ascending tract has three sets of neurons: the first order, the second order, and the third
order of neurons.
 First Order of Neurons: These are primary afferent neurons that receive sensory
impulses from the receptors and send them to sensory neurons present in the posterior
gray horn of spinal cord through their fibers. Nerve cell bodies of these neurons are
located in the posterior dorsal root ganglion. After entering the spinal cord, they
terminate on the second order of neurons in the spinal cord laminae itself or ascend up in
dorsal column to terminate on the second order of neurons in the medulla.
 Second Order of Neurons: They are either located in the spinal cord or in the
brainstem. They carry sensory impulses from spinal cord to different brain areas below
cerebral cortex (subcortical areas) such as thalamus. They are sensory neurons present
in the posterior gray horn. All the ascending tracts are formed by fibers of second order
neurons of the sensory pathways except the ascending tracts in the posterior white
funiculus, which are formed by the fibers of first order neurons.
 Third Order of Neurons. These neurons originate from the specific nuclei in the
thalamus and terminate in the specific areas in the sensory cortex. Third order neurons
are in the subcortical areas that carries sensory impulses from subcortical areas to
cerebral cortex. Sensations transmitted from different parts of body are relayed in the
thalamus before being projected into the cortex. Therefore, thalamus is considered as the
main sensory relay center in the brain.

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 Higher (Fourth) Order of Neurons: These neurons are usually not included in the
sensory pathways. However, sometimes a higher order of neuron originates from the
terminals of third order of neurons in the sensory cortex and project to other areas of the
brain, especially to the cortical sensory association area, for further processing of the
sensory information.

Classification of Ascending Tracts


The ascending pathways are typically divided into three categories: dorsal column
pathways, anterolateral systems and other ascending pathways.
 Dorsal Column Pathways: Fibers ascending in the dorsal column of the spinal cord are
included in this pathway. This is also called lemniscal system as fibers occupy medial
lemniscus in the brainstem. Dorsal column pathways transmit following sensations:
1. Fine touch
2. Vibration
3. Proprioception
4. Tactile localization
5. Tactile discrimination
6. Stereognosis
 Anterolateral Systems: It is divided into two parts: anterior spinothalamic tract and
lateral spinothalamic tract.
1. The anterior spinothalamic tract carries the sensation of crude touch.
2. The lateral spinothalamic tract carries the sensation of pain and temperature.
 Other Ascending Pathways: These include many other ascending pathways such as:
1. Spinocerebellar tract (dorsal and ventral)
2. Spinoreticulothalamic tract
3. Spinocervicothalamic tract

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A. Dorsal column pathways. B. Anterolateral system.

a. Dorsal Column Pathways


The large myelinated afferent fibers that enter the dorsal column of the spinal cord to
reach the medulla constitute dorsal column pathways. They carry the sensations of fine touch,
vibration, proprioception, tactile localization, tactile discrimination and stereognosis. In the

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spinal cord, they ascend up in two fasciculi: the gracile fasciculus and the cuneate fasciculus.
These ascending tracts in spinal cord are also called tract of Goll and Burdach.

First Order of Neuron: Neurons arriving from lower extremity and lower part of the trunk
ascend up in the gracile fasciculus, whereas the neurons arriving from upper extremity and upper
part of the trunk ascend in the cuneate fasciculus.
1. The gracile fasciculus is located medially in the spinal cord and carries sensations from
hindlimb and trunk.
2. The cuneate fasciculus is situated laterally that transmits impulses from upper limb and
upper part of the trunk.
3. The first order of neurons terminate in the nucleus gracilis and nucleus cuneatus in the
medulla.

Second Order of Neuron: The cell bodies of these neurons are present in the nucleus gracilis
and cuneatus in the medulla.
1. These two nuclei are known as the dorsal column nuclei (cluster andnon-cluster zones).
2. The fibers originating from these nuclei cross the midline and pass on to the opposite side in
the medulla and ascend up in the medial lemniscus to reach thalamus.
3. The second order of neurons, thus transmit impulses to the contralateral thalamus.

Third Order of Neuron: These neurons originate from the specific nucleus (the ventral
posterior and lateral nucleus) in the thalamus. The neurons project to the somatosensory areas
of the cerebral cortex.

b. Anterolateral System
Phylogenetically, this is older than the dorsal column system. It is divided into anterior
spinothalamic tract that carries the sensation of crude touch, and lateral spinothalamic tract
that carries the sensation of pain and temperature
First Order of Neuron: These are primarily the afferent fibers originating from
nociceptors, thermoreceptors, and mechanoreceptors.
1. The fibers enter the spinal cord through dorsal root and the cell bodies are present in the DRG.

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2. In the spinal cord, fibers terminate on the second order of neurons that are present on the
same side of the dorsal horn of the spinal cord.
3. The cell bodies of second order of neurons are located mainly in the laminae I, II and V in the
dorsal horn.

Second Order of Neuron: The cell bodies of these neurons are present in the dorsal horn of the
spinal cord
1. The axons cross the midline in the same spinal segment and ascend up in the opposite side of
the anterolateral funiculus to reach the thalamus.
2. The fibers carrying the sensation of crude touch are placed anteriorly and therefore called
anterior or ventral spinothalamic tract.
3. The fibers carrying the sensations of pain and temperature are placed laterally and therefore
called lateral spinothalamic tract.

Third Order of Neuron: The neurons originate from the VPL, midline and intralaminar nuclei
of the thalamus and project to the specific areas in the sensory cortex.
Table 1: List of ascending tracts of spinal cord
White Column Tract
Anterior white column Anterior spinothalamic tract
Lateral white column Lateral spinothalamic tract
Ventral spinocerebellar tract
Dorsal spinocerebellar tract
Spinotectal tract
Fasiculus dorsolateralis
Spinoreticular tract
Spino-olivary tract
Spinovestibular tract
Posterior white Fasciculus gracilis
Column Fasciculus cuneatus
Comma tract of Schultze

DESCENDING TRACTS OF SPINAL CORD


Descending tracts of the spinal cord are formed by motor nerve fibers arising from brain
and descend into the spinal cord. Descending pathways are fibers passing downward from
different parts of the brain that influence spinal cord neurons. However, all descending tracts are
not motor pathways like ceruleospinal and raphespinal tracts that predominantly influence

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sensory functions. That is to say that Supraspinal influences on motor control are mediated
through descending pathways.
Supraspinal centers controlling motor activities are broadly divided into two categories:
the motor cortex and the brain stem centers. Motor cortex influences spinal motor neurons via
corticospinal tracts and brainstem motor nuclei influence motor functions mainly via
rubrospinal, reticulospinal and vestibulospinal tracts.

Classification of Descending Tracts


Descending motor pathways have been traditionally classified into two types: Pyramidal
tracts and extrapyramidal tracts.

Pyramidal Tracts
Pyramidal tracts were the first tracts to be found in man. Pyramidal tracts of spinal cord
are the descending tracts concerned with voluntary motor activities of the body. These tracts are
otherwise known as corticospinal tracts. There are two corticospinal tracts, the anterior
corticospinal tract and lateral corticospinal tract. While running from cerebral cortex towards
spinal cord, the fibers of these two tracts give the appearance of a pyramid on the upper part of
anterior surface of medulla oblongata hence the name pyramidal tracts

Extrapyramidal Tracts
Descending tracts of spinal cord other than pyramidal tracts are called extrapyramidal
tracts.
Table 2: List of descending tracts of spinal cord
Type Tract
Pyramidal tracts Anterior corticospinal tract
Lateral corticospinal tract
Extrapyramidal tracts Medial longitudinal fasciculus
Anterior vestibulospinal tract
Lateral vestibulospinal tract
Reticulospinal tract
Tectospinal tract
Rubrospinal tract
Olivospinal tract

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