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1
i. Abstract
ii. Content
Part I: Proprioception
1. Introduction to proprioception
2. Proprioceptors
5. Conscious proprioception
11. Summary
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iii. Checklist of topics and activities in this
module
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Activity 15: Control of posture and movement at the
brain stem level
9 Control of posture and movement at the
cerebral cortex level
Activity 16. Cortical motor areas
Activity 17. Motor homunculus
10 Control of posture and movement at the
associate areas level
ix Summary
x Conclusion
Web Animation:
v. Background knowledge
To complete this module successfully, you should have the following background:
• The anatomy and histology of the nervous system (relevant General Anatomy
modules).
• The whole Introduction to Medical Physiology modules
Please add other terms that you feel are relevant to your understanding of this
module.
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vii. Objectives
After studying the materials in this module, the students should be able to:
Please set up more specific objectives after you have thoroughly studied the
material in this module to help yourself in your revision later on. Write notes
that meet the requirement of the new objectives.
NEU 88. Draw a cross section of the spinal cord and discuss the organization of the sensory and motor
components of gray matter. Describe the somatotopic arrangement of motor neuron pools.
NEU 89. List the medial and lateral motor systems. Describe their origin, pathway, and termination within
the spinal cord. Compare their functions in motor control.
NEU 90. Describe the effects of lesions in medial and lateral systems.
NEU 92. List three functional divisions of the cerebellum, detailing the input and output connections of each.
Be able to differentiate the functions of each and their integration with lateral and medial motor systems.
NEU 93. Draw and label the circuitry of the cerebellar cortex, assign the functional role of each neuron type
and give its synaptic action (excitatory/inhibitory). Be able to describe how this circuit functions as a timing
mechanism and how it produces synergy in opposing muscle groups.
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NEU 94. On the basis of input-output organization, somatotopic organization, and overall function, predict
the neurological disturbances that can result from disease or damage in different regions of the cerebellum.
NEU 95. Contrast the spinal proprioceptive pathways to the cerebellum with those to the cortex.
NEU 96. List and describe the major interconnections between components of the basal ganglia and the
motor cortex. Identify the neurotransmitters determining the flow of information in the system.
NEU 97. Describe the overall function of the basal ganglia in movement control and initiation in association
with medial and lateral motor systems.
NEU 98. List the appropriate signs of rigidity, dyskinesias, akinesia, and tremor for Parkinsonism, chorea,
hemiballism, and athetosis. Assign a likely lesion site or chemical system defect for each clinical syndrome.
NEU 99. Describe the rationale for treatment of Parkinsonism with anticholinergic drugs, L-DOPA, or
transplantation of catecholamine-producing cells.
Cerebral Cortex
NEU 100. Describe the medial to lateral, rostral to caudal, and surface to white matter organizations of the
primary motor cortex and the premotor cortex. Draw those regions on a sketch of the brain and also locate
the supplementary motor cortex.
NEU 101. Compare the effects of electrical stimulation of motor cortex and premotor cortex, relating the
expected results to the control of voluntary movement.
NEU 102. Describe the origin, course, and termination of the pyramidal tract.
NEU 103. Compare the consequences of upper motor neuron loss to lower motor neuron loss. Describe the
consequences of pyramidal tract transection.
NEU 104. Draw a “flow diagram” for the brain regions involved in planning, initiating, and properly executing
a skilled voluntary movement.
NEU 105. Identify Brodmann areas for visual, auditory, somatic sensory, motor, and speech areas.
NEU 106. Identify the cortical areas that receive projections from the following thalamic nuclei: ventral
lateral, dorsomedial, pulvinar, medial geniculate, lateral geniculate, ventral posterolateral, and posteromedial.
NEU 108. Describe the cortical area important for spatial relations.
NEU 110. Define and explain the physiological basis of evoked potentials and the electroencephalogram
(EEG). List the main clinical uses of each.
NEU 111. Describe the primary types of rhythms that make up the EEG and the behavioral states that
correlate with each.
NEU 112. Describe the origin of spontaneous electrical activity of the cerebral cortex.
NEU 113. Distinguish EEG activity from evoked potentials and the uses of evoked potentials.
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viii. Learning Activities
Part I: Proprioception
1. Introduction to proprioception
Definition:
Proprioception= sense of position of body parts
Kinesthesia = sense of movement of body parts
In Module 18 we discussed the flow of information from receptor to control centre to effector in
accomplishing various reflexes. In such cases, the stimuli could come from the internal or external
environment (please give examples), but the control centre does not involve the conscious part of
the brain (where are the control centres for reflexes?). We have also previously agreed that
external environmental stimuli are perceived consciously (sight, smell, hearing, taste, touch) and
internal environmental stimuli (blood pressure, PO2, etc) are not.
In contrast, the sense of position of our body parts (proprioception) could be consciously
perceived or nonconsciously sensed. It is consciously perceived when we put our thoughts into it
but normally we are not conscious about the positions of our body parts. For example, we know
where our limbs are even with our eyes are closed and we can control their movement consciously
when we think about them, but they normally assume positions (for appropriate postures or even
for the act of walking) without much conscious control at all. Regardless of whether we are
conscious of the positions and control of our body parts or not, afferent information about them is
being sent to the control centre all the time (mostly to the subcortical areas), and efferent
information is continually sent to the effectors to adjust our posture appropriately. This is how we
assume our minute-to-minute positions of our body.
How is information that brings about proprioception generated? How is the information processed?
What is the significance of the sensory information? How do we go about controlling our minute-
to-minute posture? What is the nature of the efferent information? We’ll try to address these issues
in this module.
Proprioception provides feedback solely on the status of the body internally. It is the sense that
indicates whether the body is moving with required effort, as well as where the various parts of the
body are located in relation to each other. Thus, proprioception correctly describes afferent
information arising from internal peripheral areas of the body that contribute to postural control,
joint stability, and several conscious sensations. Proprioceptive information concerning the status
of the joint and associated structures is essential for neuromuscular control. Proprioception is
conveyed to all levels of the central nervous system, where it provides a unique sensory component
to optimize motor control.
This module will be basically divided into 2 main parts: 1. Proprioception, and 2. Posture control.
We’ll focus on the afferent segment of proprioception and the efferent segment of posture control.
We’ll start off with how afferent inputs are generated by the proprioceptors for perception to take
place.
• What is the significance of the phenomena that external environmental stimuli are perceived
consciously (sight, smell, hearing, taste, touch) and internal environmental stimuli (blood
pressure, PO2, etc) are not?
• What are proprioceptors? Give examples. What would stimulate proprioceptors? What is the
significance of proprioception?
• What happens to the afferent information coming from proprioceptors? What is the significance
that most of the afferent information coming from proprioceptors are not consciously perceived?
• What happens to the efferent information resulting from integration of afferent information coming
from proprioceptors?
• What are the effectors for controlling body posture and movement? Why is body posture
important? Explain how it is controlled consciously and nonconsciously.
• Draw a model that shows the relationship between proprioceptors, control centres and effectors.
Identify the control centres and effectors. Identify the ascending and descending pathways in the
model.
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2. Proprioceptors
The awareness of the orientation of the body in space (proprioception) and the direction, extent,
and rate of movement of the limbs (kinesthesia) depend in part upon information derived from
sensory receptors in the joints (joint receptors, Fig. 2), tendons (Golgi tendon organ, Fig 4), and
muscles (muscle spindle, Fig 3). Information from these proprioceptors is normally integrated
with that arising from vestibular receptors (which signal gravitational acceleration and changes in
velocity of movements of the head), as well as from visual, auditory, and tactile receptors.
Sensory information from certain proprioceptors, particularly those in muscles and tendons, need
not reach consciousness, but can be used by the motor system as feedback to guide postural
adjustments and control of well-practiced or semiautomatic movements such as those involved in
walking.
Although visual and vestibular input contributes, the peripheral mechanoreceptors are the most
important from a clinical orthopaedic perspective (Fig. 1). Afferent pathways (dotted lines) convey
input to the 3 levels of motor control areas (spinal cord, brain stem and cerebral cortex) and
associated areas such as the cerebellum. Activation of motor neurons may occur in direct response
to peripheral sensory input (reflexes) or from descending motor commands, both of which may be
modulated or regulated by the associate areas (gray lines). Efferent pathways from each of the
motor control levels (solid lines) converge upon the alpha and gamma motor neurons located in the
ventral aspects of the spinal cord. The contractions by the extrafusal and intrafusal muscle fibers
cause new stimuli to be presented to the peripheral mechanoreceptors.
• Where do afferent
information for
proprioception come
from? Name the
proprioceptors. What is
the information from
these proprioceptors
integrated with?
In summary, information for proprioception and kinesthesia is derived from proprioceptors, and is
normally integrated with that arising from
a. vestibular receptors (which signal gravitational acceleration and changes in velocity
of movements of the head),
b. visual, auditory, and tactile receptors
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Activity 3. Joint receptors
The joint 1° afferents respond to changes in the angle, direction, and velocity of movement in a
joint. The responses are predominantly rapidly adapting with few joint 1° afferents signaling the
resting (static) position of the joint. It has been suggested that information from muscles, tendons,
skin and joints are combined to provide estimates of joint position and movement. For example,
when the hip joint is replaced — removing all joint receptors — the ability to detect the position of
the thigh relative to the pelvis is not lost.
Although 4 types of receptors are dispersed throughout ligamentous and capsular tissues, Ruffini
receptors are the most frequently described. They are considered to behave as both static and
dynamic receptors based on their low-threshold, slow-adapting characteristics. In contrast, the low-
threshold, rapidly adapting characteristics of Pacinian corpuscles cause them to be exclusively
classified as dynamic receptors. Also present in these tissues are Golgi tendon organ-like endings
and free nerve endings.
Golgi tendon organs (GTOs, Fig. 4) are mechanoreceptors located within musculotendinous tissue
Through each GTO passes a small bundle of muscle tendon fibers destined to attach to muscle
fibers. This series arrangement, coupled with the very low threshold and high dynamic sensitivity
exhibited by the sensory endings, enables GTOs to provide the CNS with feedback concerning
muscle tension. GTOs function primarily in signaling active muscle tension (tension developed
during contraction) rather than passive tension (tension developed during inactive muscle
stretching).
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Muscle spindles (Fig. 3). As a whole, muscle spindles are responsible for conveying information
regarding muscle length and rate of changes in length. Muscle spindles consist of specialized
afferent nerve endings that are wrapped around modified muscle fibers (intrafusal fibers), several of
which are enclosed in a connective tissue capsule. There are different types of intrafusal fibers:
some are mainly sensitive to changes in muscle length, whereas others are more sensitive to the rate
of change in muscle length.
Although the central areas of the intrafusal muscle fibers lack contractile elements, the peripheral
areas contain contractile elements, which are innervated independent of extrafusal (skeletal) muscle
fibers via the gamma motor neurons (γ MNs). Activation of the peripheral contractile elements
stretches the central regions containing the sensory receptors from both ends. This results in an
increase in the firing rates of the sensory ending and an increase in the sensitivity of the muscle
spindle to length changes. At the spinal level, various peripheral receptors, such as skin receptors,
articular receptors, and chemoreceptors, strongly influence the activity of the γ-MN system and,
therefore, the muscle spindle in providing afferent information.
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Activity 4: Proprioceptors
• Draw diagrams to depict proprioceptors in the muscle, tendon and joints. Describe how these
receptors are activated.
• Describe how afferent information is carried to the CNS. Discuss how it affects motor control at
the level of the spinal cord, brain stem and cerebral cortex.
• Explain how information from other receptors (special senses, tactile senses) affects afferent
and efferent information with regard to postural control.
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3. Sensory integration at the spinal cord level
Many of the axons conveying proprioceptive information bifurcate once they enter the dorsal horn
of the spinal cord to synapse with interneurons. The essence of afferent integration at the spinal
cord level lies with the interneurons and the neurons connecting with higher CNS levels. Control
over these neurons via descending commands from the brain stem and cortex provides these centers
with the ability to filter the sensory input that will be conveyed via the ascending tracts. In other
words, the supraspinal CNS regions modulate the sensory information from the periphery that
enters the ascending tracts.
Integration at the spinal cord level is also influenced by the final common input to the α-motor
neuron (Fig. 3). This hypothesis resides on the strong influence that the muscle, skin, and joint
afferents and descending pathways have over gamma neuron activation. As mentioned previously,
the peripheral regions of intrafusal muscle fibers contain contractile elements innervated by γ MNs,
with the level of activation directly controlling muscle spindle sensitivity. Any of the signals
barraging the γ-MN pools alter their level of activation, and, therefore, influence the input arising
from the muscle spindles. Thus, the afferent signals from muscle spindles are hypothesized to be a
function of muscle length changes superimposed on the integrated peripheral receptor and
descending pathway information. In this manner, the γ-MN system may be considered a “premotor
neuronal integrative system” that conducts “polymodal feedback” to the CNS (Fig. 3).
Activity 5: Sensory
ntegration at spinal cord
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4. Ascending spinal tracts conveying proprioceptive information
Most proprioceptive information travels to higher CNS levels through either the dorsal lateral
tracts or the spinocerebellar tracts. The 2 dorsal lateral tracts are located in the posterior region of
the spinal cord and ultimately convey the signals to the somatosensory cortex. Although the
majority of the sensations traveling in this tract are touch, pressure, and vibration, various amounts
of the conscious appreciation of position and kinesthetic sensations have also been attributed to this
tract.
The spinocerebellar tracts are characterized by the fastest transmission velocities in the CNS. As
their name suggests, the spinocerebellar tracts terminate in various areas of the cerebellum, where
the signals may be processed and integrated with other afferent and descending information. In
contrast to the conscious sensory appreciation associated with the dorsal lateral tracts, the
spinocerebellar tracts are believed to be responsible for “nonconscious proprioception” (i.e. limb
position, joint angles, and muscle tension and length) used for reflexive, automatic, and voluntary
activities. In addition to relaying peripheral afferent information, parts of these tracts are associated
with transmitting an efferent copy of motor neuron drive back to the higher CNS levels.
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5. Conscious proprioception
As mentioned previously, proprioception information for conscious appreciation travels via the
dorsal lateral tracts, but the contributions to these tracts from muscle and joint mechanoreceptors
remaining largely unknown. Thus, projections to the cortical sensory areas and conscious
perceptions after direct receptor stimulation is the second necessity in determining the predominant
source of conscious proprioception (Figure 4). Cortical projections have been reported from joint
(both capsular and ligament) afferents, muscle spindles, and GTOs. It has been demonstrated that
mechanical stimulation of cutaneous receptors elicited kinesthetic sensations. While direct
stimulation of a single muscle spindle afferent failed to elicit movement perception, stimulation of
several muscle spindles through vibration and isolated traction has been reported to evoke
conscious movement sensations. The failure of joint and cutaneous afferents anesthesia to disrupt
conscious kinesthesia and JPS provides further support for the importance of muscle receptors in
conscious proprioception.
Activity 7. Conscious
proprioception
In summary, proprioception is the awareness of the orientation of the body in space. The afferent
information from proprioceptors is integrated with information from vestibular receptors (which
signal gravitational acceleration and changes in velocity of movements of the head), as well as from
visual, auditory, and tactile receptors). This could be consciously perceived or non-consciously
processed to produce appropriate motor functions i.e. posture control.
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Part II: Control of Posture
So far we have discussed the afferent segment involved in proprioception. The significance of
proprioception is so that we are able to adjust body posture and body movement appropriately,
either voluntarily or involuntarily or both. We’ll now look at the efferent segment i.e. how our
movement and body positions are adjusted to suit the activities that we do.
Posture may be defined as the position adopted by the individual within his or her environment.
Muscles that help us manage our posture are mostly axial and antigravity muscles. These muscles,
like most skeletal muscles are always in a partial state of contraction, so they exhibit some degree
of tone. We are able to stand and sit comfortably for hours without feeling fatigue, because of these
muscles, in particular their tone. The tone of these muscle and the maintenance of an upright
posture, in turn involves postural reflexes which include the stretch reflex and the crossed-
extensor reflex which have already been discussed (Module 18).
Along with the maintenance of upright posture is the maintenance of balance which is a complex
process in the human because of our tall height, which must be balanced over the feet which have a
small area. Adding to the difficulty of maintaining balance and posture is our high center of gravity,
centered over the hips.
• What is posture? What is postural reflex? Hint: relate the relevant receptor to control centre
to effector.
• Compare and contrast between axial muscles and distal muscles. Which motor pathways is
their contraction mediated by? Give examples.
• Explain how stretch reflex and cross extensor reflex are involved in posture regulation.
• What are the organs that have receptors that provide sensory inputs for postural reflexes?
The postural reflexes are aided by afferent sensory information from the eyes, the vestibular
apparatus, and the somatic receptors (proprioceptors) and the efferent response is to the skeletal
muscles after integration in the brainstem and spinal cord. Control of posture involves active
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muscular resistance to displacement of the body. The crossed-extensor reflex is one example of a
postural reflex. As one leg is flexed, the other is extended to support the added weight of the body.
In addition, the positions of various parts of the body are shifted to move the center of gravity over
the single, weight-bearing leg. This shift in the center of gravity is important in the stepping
mechanism of walking.
The motor components of the sensorimotor system contributing to posture consist of a central axis
and 2 associate areas. The central axis corresponds to the 3 levels of motor control, spinal cord,
brain stem, and cerebral cortex, whereas the 2 associate areas, cerebellum and basal ganglia, are
responsible for modulating and regulating the motor commands. Sensory information underlies the
planning of all motor output and is conveyed to all 3 levels of motor control.
Activation of motor neurons may occur in direct response to peripheral sensory input (reflexes) or
from descending commands initiated in the brain stem or cerebral cortex, or both. Independent of
the initiating source, skeletal muscle activation occurs through signal convergence onto the motor
neurons located in the spinal ventral horns i.e. the final common path (Module 13). Both types of
motor neurons, alpha motor neurons (α MNs) controlling extrafusal muscle fibers (skeletal) and γ
MNs controlling intrafusal muscle fibers (muscle spindles), exit the spinal ventral horns.
The central axis areas are organized in both a hierarchic and parallel manner (Module 17). The
hierarchic organization allows the lower motor areas to automatically control the details of common
motor activities, while the higher centers can devote resources to controlling the more precise and
dexterous motor activities. In addition, as mentioned earlier, higher levels can regulate the afferent
information reaching them through inhibitory and facilitatory control over sensory relay nuclei.
Through the parallel arrangement, each motor control center can directly issue independent
contributory descending motor commands directly on the motor neurons.
The final common paths to skeletal muscle (the spinal motor neurons and homologous neurons in
the motor nuclei of the cranial nerves), are bombarded by impulses from an immense array of
pathways:
a. Each spinal motor neuron has many inputs from the same spinal segment (e.g: reflexes).
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b. Numerous supra-segmental inputs also converge on these cells from other spinal segments, the
brainstem, and the cerebral cortex.
Some of these inputs end directly on the motor neurons, but many exert their effects via
interneurons or via the -efferent system to the muscle spindles and back through the Ia afferent
fibers to the spinal cord.
• Draw an appropriate diagram to show the neuronal connections to the motor neuron.
It is the integrated activity of these multiple inputs from spinal, medullary, midbrain, and cortical
levels that regulates the posture of the body and makes coordinated movement possible.
The inputs converging on the motor neurons subserve three semi-distinct functions:
• they bring about voluntary activity;
• they adjust body posture to provide a stable background for movement; and
• they coordinate the action of the various muscles to make movements smooth and precise.
In summary, the control of posture and movement involve the following activities:
• The patterns of voluntary activity are planned within the brain.
• The commands are sent to the muscles primarily via the corticospinal and corticobulbar
systems.
• Posture is continually adjusted not only before but also during movement by posture-
regulating systems.
• Movement is smoothed and coordinated by the medial and intermediate portions of the
cerebellum (spinocerebellum) and its connections.
• The basal ganglia and the lateral portions of the cerebellum (neocerebellum) are part of a
feedback circuit to the premotor and motor cortex that is concerned with planning and
organizing voluntary movement.
Motor output is of two types: reflexive or involuntary, and voluntary. Some would add as a
subdivision of reflex responses rhythmic responses such as swallowing, chewing, scratching, and
walking, which are largely involuntary but subject to voluntary adjustment and control.
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The motor system "learns by doing", and performance improves with repetition. This involves
synaptic plasticity (the ability of the synapse between two neurons to change in strength).
Commands for voluntary movement originate in cortical association areas (Fig. 9). The movements
are planned in this cortical area as well as in the basal ganglia and the lateral portions of the
cerebellar hemispheres. The basal ganglia and cerebellum both funnel information to the premotor
and motor cortex by way of the thalamus. Motor commands from the motor cortex are relayed in
large part via the corticospinal tracts to the spinal cord and the corresponding corticobulbar tracts to
motor neurons in the brainstem. However, collaterals from these pathways and a few direct
connections from the motor cortex end on brainstem nuclei, which also project to motor neurons in
the brainstem and spinal cord. These pathways can also mediate voluntary movement.
Movement sets up alterations in sensory input from the special senses and from muscles, tendons,
joints, and the skin. This feedback information, which adjusts and smoothes movement, is relayed
directly to the motor cortex and to the spinocerebellum. The spinocerebellum projects in turn to the
brainstem. The main brainstem pathways that are concerned with posture and coordination are the
rubrospinal, reticulospinal, tectospinal, and vestibulospinal tracts and corresponding projections to
motor neurons in the brainstem.
A. Rubrospinal Tract: Originates in the red nucleus. Fibers project to interneurons in the
spinal cord which excite motoneurons of flexor muscles and inhibit motoneurons of
extensor muscles to release the body from a postural stance.
D. Lateral Vestibulospinal Tract: This tract originates in the lateral vestibular nucleus
(Dieter's nucleus) and projects to motoneurons and interneurons. Stimulation of cells in the
nucleus produces a powerful excitation of extensors and inhibition of flexors. This tract
plays an important role in the control of antigravity muscles and the maintenance of posture.
With the aid of a diagram, discuss the sequence of events that take place for movement to
occur i.e. starting from sensory input.
The nerve fibers that pass from the motor cortex to the cranial nerve nuclei form the corticobulbar
tract. The nerve fibers that cross the midline in the medullary pyramids and form the lateral
corticospinal tract make up about 80% of the fibers in the corticospinal pathway. The remaining
20% make up the anterior, or ventral, corticospinal tract (Fig. 10), which does not cross the
midline until it reaches the level of the muscles it controls. At this point, its fibers end on
interneurons that make contact with motor nerves on both sides of the body. The lateral
corticospinal tract is concerned with skilled movements, and in humans its fibers end directly on
the motor neurons.
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6.4. Control of axial and distal muscles
Another important theme in motor control is that in the brainstem and spinal cord,
• medial or ventral pathways and neurons are concerned with the control of muscles of the
trunk and proximal portions of the limbs
• lateral pathways are concerned with the control of muscles in the distal portions of the limbs.
The axial muscles are concerned with postural adjustments and gross movements, whereas the
distal limb muscles are those that mediate fine, skilled movements. Thus, for example, the neurons
in the medial portion of the ventral horn innervate the proximal limb muscles, particularly the
flexors, whereas the lateral ventral horn neurons innervate the distal limb muscles. Similarly, the
ventral corticospinal tract and the medial descending paths from the brainstem (the tectospinal,
reticulospinal, and vestibulospinal tracts) are concerned with adjustments of proximal muscles and
posture, whereas the lateral corticospinal tract and the rubrospinal tract are concerned with distal
limb muscles and, particularly in the case of the lateral corticospinal tract, with skilled voluntary
movements.
Draw a cross section of a spinal cord showing the positions of sensory and motor tracts. Identify the
tracts that contain nerve fibres responsible for movements of axial muscles and distal limb muscles.
It should be apparent from our earlier discussion that the spinal cord plays an integral role in motor
control, despite the gross anatomy suggesting it may only be a medley of conduction pathways.
From the spinal cord arise direct motor responses to peripheral sensory information (reflexes) and
elementary patterns of motor coordination (rhythmic and central pattern generators). As discussed
earlier, very little afferent input and few descending commands synapse directly on motor neurons.
Instead, most input terminates upon the interneurons located throughout all areas of cord gray
matter. Even in the case of a simple monosynaptic reflex, such as the stretch reflex, birfurcations
from the incoming afferent fiber arise. These bifurcations may convey the afferent information to a
number of locations, including interneurons, higher motor centers, and other motor neurons
(antagonistic). The bifurcations and interneuronal networks provide the basis for the spinal cord's
efferent integrative functions.
Reflexes may be elicited from the stimulation of cutaneous, muscle, and joint mechanoreceptors
and may involve excitation of α-MNs, γ-MNs, or both. For many clinicians, the stretch reflex in
response to rapid muscle lengthening provides the most familiar example. These reflexes, as well as
the other reflexes attributed to the spinal cord neuronal circuitry, are more complex than simple
direct input-output connections. Superimposed on even the simplest monosynaptic reflexes are
influences from such sources as other afferent input, descending commands, or both.
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Activity 14: Control of posture and movement at the spinal cord level
• Draw a cross section of the spinal cord. Indicate the grey matter and the white matter.
• Draw the incoming afferent fibre and outgoing lower motor neuron.
• Draw the synapses on the lower motor neuron i.e. directly from the afferent fibre and the
descending fibes; indirectly via interneurons. Discuss the significance of the spinal interneurons.
• Summarise how the spinal cord is involved in the control of posture and movement.
Despite being the most primitive part of the brain from a phylogenetic perspective, the brain stem
contains major circuits that control postural equilibrium and many of the automatic and stereotyped
movements of the body. In addition to being under direct cortical command and providing an
indirect relay station from the cortex to the spinal cord, areas of the brain stem directly regulate and
modulate motor activities based on the integration of sensory information from visual, vestibular,
and somatosensory sources.
Two main descending pathways, the medial and lateral pathways, extend from the brain stem to
the spinal cord neural networks. The medial pathways influence the motor neurons innervating the
axial and proximal muscles, while the lateral pathway controls the distal muscles of the extremities.
In addition to controlling postural control, some axons comprising the medial pathways make
excitatory and inhibitory (including suppression of spinal reflexes) synapses with the interneurons
and motor neurons involved with movement and postural control. Through influences on the γ-
MNs, parts of both the medial and lateral tracts assist in maintaining and modulating muscle tone.
Activity 15: Control of posture and movement at the brain stem level
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9. Control of posture and movement at the cerebral cortex level
In general, the motor cortex is responsible for initiating and controlling more complex and discrete
voluntary movements. It is divided into 3 specialized and somatotopically organized areas, each of
which project directly and indirectly (via the brain stem) onto interneurons and motor neurons
located in the spinal cord.
1. the primary motor cortex, receives peripheral afferent information via several pathways
and is responsible for encoding the muscles to be activated, the force the recruited muscles
produce, and the direction of the movement.
2. the premotor area, also receives considerable sensory input; however, it is mainly involved
with the organization and preparation of motor commands.
3. the supplemental motor area also plays an important role in programming complex
sequences of movement that involve groups of muscles.
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The various parts of the body are represented in the precentral gyrus, with the feet at the top of the
gyrus and the face at the bottom (Fig. 4). The facial area is represented bilaterally, but the rest of
the representation is generally unilateral, the cortical motor area controlling the musculature on the
opposite side of the body. The cortical representation of each body part is proportionate in size to
the skill with which the part is used in fine, voluntary movement. The areas involved in speech and
hand movements are especially large in the cortex; use of the pharynx, lips, and tongue to form
words and of the fingers and apposable thumbs to manipulate the environment are activities in
which humans are especially skilled.
The major direct descending pathway from the motor cortex to the α MNs and γ MNs is the
corticospinal tract. In addition to influencing motor functions directly, the corticospinal tract also
affects motor activity indirectly through the descending brain stem pathways.
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10. Control of posture and movement at the associate areas level
Although the 2 associate areas, the cerebellum and basal ganglia, cannot independently initiate
motor activity, they are essential for the execution of coordinated motor control. The cerebellum,
operating entirely at a subconscious level, plays a major role in both the planning and modification
of motor activities though comparison of the intended movement with the outcome movement.
This is accomplished through the continuous inflow of information from the motor control areas
and the central and peripheral sensory areas. The cerebellum is divided into 3 functional divisions.
The first division receives vestibular input, both directly and indirectly from the vestibular labyrinth
(semicircular and otolith receptors) and, as might be surmised based on the input, is involved with
postural equilibrium. The second cerebellar division is mainly responsible for the planning and
initiation of movements, especially those requiring precise and rapid dexterous limb movements.
This division receives input from both the sensory and motor cortices. It is the third division, the
spinocerebellum, which receives the somatosensory information conveyed through the 4 ascending
spinocerebellar tracts. In addition to the somatosensory input, this division of the cerebellum also
receives input from the vestibular labyrinth and visual and auditory organs. The output from the
spinocerebellum serves to adjust ongoing movements through influential connections on the medial
and lateral descending tracts in the brain stem and cortex via projections on the vestibular nucleus,
reticular formation, red nucleus, and motor cortex. In addition to controlling movements, the
spinocerebellum also uses the somatosensory input for feedback regulation of muscle tone through
regulation of static γ-MN drive to the muscle spindles. Lastly, the cerebellum also receives an
efferent copy of the motor commands arriving at the ventral roots of the spinal cord. The
cerebellum has also been implicated in motor learning.
The basal ganglia consist of 5 subcortical nuclei (groups of nerve cells) located deep within the
cerebral hemispheres. In contrast to the cerebellum, which has input and output connections with
all 3 levels of motor control, the cerebral cortex is the only central axis component having input and
output connections (via the thalamus) with the basal ganglia. With respect to motor control, the
basal ganglia are believed to be involved with more higher-order, cognitive aspects of motor
control. An additional distinction from the cerebellum is that the basal ganglia receive input from
the entire cerebral cortex, not just those associated with sensory and motor function. The
widespread input and output cortical connections suggest that they are involved with many
functions other than motor control.
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ix. Summary
It is impossible to separate postural adjustments from voluntary movement in any rigid way, but it
is possible to differentiate a series of postural reflexes (Table 2) that not only maintain the body in
an upright, balanced position but also provide the constant adjustments necessary to maintain a
stable postural background for voluntary activity. These adjustments include maintained static
reflexes and dynamic, short-term phasic reflexes. The former involve sustained contraction of the
musculature, whereas the latter involve transient movements. Both are integrated at various levels
in the CNS from the spinal cord to the cerebral cortex and are effected largely through various
motor pathways. A major factor in postural control is variation in the threshold of the spinal stretch
reflexes, which is caused in turn by changes in the excitability of motor neurons and, indirectly, by
changes in the rate of discharge in the efferent neurons to muscle spindles.
Table 2. Postural reflexes
Reflex Stimulus Response Receptor Integrated
In
Stretch reflexes Stretch Contraction of muscle Muscle spindles Spinal cord,
medulla
Positive supporting Contact with sole or palm Foot extended to support Proprioceptors in Spinal cord
(magnet) reaction body distal flexors
Neck righting Stretch of neck muscles Righting of thorax and Muscle spindles Midbrain
reflexes shoulders, then pelvis
Body on body Pressure on side of body Righting of body even Exteroceptors Midbrain
righting reflexes when head held sideways
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Hopping reactions Lateral displacement while Hops, maintaining limbs in Muscle spindles Cerebral
standing position to support body cortex
Transections at different levels of the brainstem have been used to demonstrate the importance of
brainstem centers in the control of posture. Isolation of centers below the transection from central
influences above, reveals the regulatory functions of the intact centers.
A. Spinal Transection
2. Loss of conscious sensation below the level of the lesion. Sensory information
from the body regions below the transection cannot reach higher centers and those
regions appear to be anesthetized.
Two brainstem centers that are important for the maintenance of muscle tone in antigravity
muscles (the extensors) are the pontine reticular formation (medial reticulospinal tract)
and Dieter's nucleus (lateral vestibulospinal tract). Both centers have an excitatory
influence on extensor muscles. Stimulation of cells in the pontine reticular formation has a
very powerful excitatory effect on extensors, but its activity is normally inhibited by central
(cortical) projections. If the spinal cord is cut above the level of the pontine reticular
formation, (mid-collicular), the inhibitory influence is removed and there is an exaggerated
activation of muscle tone in the extensors (antigravity muscles). This produces a rigid
posture which is referred to as decerebrate rigidity. In patients with this condition, the
arms and legs are extended, the back is arched, the head is dorsiflexed, and the feet are
pointed with the toes curled. This stiff posture does not permit the joints to bend and the
body is capable of standing upright. This is very different from spinal transection, where
extensor muscle tone is abolished and the body becomes limp.
Interruption of the corticospinal tract (with the brainstem circuitry intact) produces
decorticate rigidity. In this condition the extensors of the legs and the flexors of the arms
contract steadily. One reason for this is that the rubrospinal tract in humans only projects as
far as the cervical cord and may counteract vestibulospinal facilitation of arm but not leg
extensors.
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ix. Conclusion
28
APPENDIX
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Complete Spinal Cord Injury: No sensory or motor function in sacral segments of spinal cord
S4-S5.
Incomplete Spinal Cord Injury: Sensory and/or motor function in sacral segments of spinal cord
S4-S5.
Paraplegia: Injury to the thoracic, lumbar or sacral level of the spinal cord, resulting in paralysis of
the lower extremities
Quadriplegia: Injuries at the cervical level of the spinal cord resulting in paralysis of the upper and
lower extremities (more commonly referred to as tetraplegia).
Sacral Sparing: Sensory or motor function at the anal mucocutaneous junction. The presence of
either is considered sacral sparing.
Spinal Shock: Often occurring soon after spinal cord injury, this is a loss of reflexes below the
level of injury with associated loss of sensorimotor functions. This condition can last for several
hours to days after initial injury.
Tetraplegia: (The more commonly used term for quadriplegia) Injury to the spinal cord in the
cervical region with resulting loss of muscle strength in all 4 extremities.
The minimal screening examination should include evaluation of the following: (1) "Normal" gait across
the room. (2) "Heel-walking" with ankles dorsiflexed. (3) "Toe-walking" on the balls of the feet with
heels elevated. (4) "Tandem" gait, in which the patient puts one foot directly in front of the other, heel
to toe, and walks an imaginary line.
Cerebellar Gait
This is a wide-based, irregular, staggering, or reeling gait, as if drunk.
Sensory-Ataxic Gait
A wide-based, short, uneven gait characterized by high steps and slapping down of the feet is seen with
proprioceptive loss, as in tabes dorsalis. The eyes may remain "glued" to the ground.
Hemiplegic Gait
With the affected spastic leg extended and internally rotated and the foot in inversion and plantar
flexion, the leg circumducts at the hip to allow the foot to clear the floor.
Paraplegic Gait
A slow, stiff shuffling gait with the toes scraping and the legs "scissoring" because of increased adductor
tone associated with spasticity is seen in myelopathy or other bilateral corticospinal tract disease.
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Dystrophic Gait
Waddling and lordotic posture may result from pelvic muscle weakness.
Parkinsonian Gait
This consists of slow starting and short shuffling steps with a tendency to accelerate ("festinate") as if
chasing the center of gravity. The posture is stooped, turns are "en bloc" with the feet moving only in
tiny steps, and there is loss of normal associated movements, such as arm swinging, that help to
maintain balance.
Apraxic Gait
Apraxia consists of an inability to execute a learned motor program. Gait apraxia is loss of the ability to
walk and results from diffuse cerebral damage—more specifically, damage to the frontal lobe—despite
normal strength and coordination. The gait is similar to a parkinsonian gait, but if severe the patient will
simply stand, partially upright, unable to "remember" how to go about walking, the feet seeming to be
"glued to the floor." Alternatively, the patient will lift and lower the feet without advancing, as if drawn to
the floor by magnetic force.
Antalgic Gait
Antalgic gait is a response to pain—favoring one leg by putting as little weight as possible on it.
Choreic Gait
Choreic gait is described as lurching, "jerky twitching," and "dancing." Falls are surprisingly rare.
enteroviruses (rare)
bulbar palsy
neurofibroma, trauma
neuropathies
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syphilis
(rare)
*Upper motor neuron disorders (eg, spinal muscular atrophies) technically involve the CNS because
the cell body of the motor neuron is located in the spinal cord.
Adapted from Tandan R, Bradley WA: Amyotrophic lateral sclerosis. Part I: Clinical features,
pathology and ethical issues in management. Annals of Neurology 18:271–280, 1985; used with
permission of Little, Brown and Company.
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