NEURAL CONTROL OF

LOCOMOTION

CONTENTS

1. INTRODUCTION

2. DEFINITION OF LOCOMOTION

3. ROLE OF DIFFERENT PARTS OF NERVOUS SYSTEM IN HUMAN

LOCOMOTION

4. THEORIES OF MOTOR CONTROL

5. CENTRAL PATTERN GENERATORS(CPGs)

6. STUDIES ON ANIMALS AND HUMAN BEINGS

7. SUMMARY

8. CONCLUSION

INTRODUCTION
 Many situations such as to escape from predators, crossing shallow rivers quite
possibly given rise to bipedilism and began what many consider to be the most significant
characteristic of hominid development : the bipedal locomotion

Locomotion is essential for survival; it allows animals to acquire food, evade

predators, or find a mate. Controlling locomotion is not an easy task. The central nervous
system (CNS) somehow must generate the locomotors pattern, generate appropriate
propulsive forces, modulate changes in the centre of gravity, coordinate multi limb
trajectories, adapt to changing conditions and changing joint positions, coordinate visual,
auditory, vestibular and peripheral afferent information, and account for the viscoelastic
properties of muscle. It must do all of this within milliseconds and usually in conjunction with
coordinating a multitude of other bodily functions and movements.

Neural control of human locomotion is not yet fully understood. There is strong
evidence which suggests that isolated neural circuits called central pattern generators exist
invertebrates to produce rhythmic oscillatory patterns, which coupled with the dynamics of
the body produce locomotion. Experiments with cats have shown that a decerebrated cat can
walk on a treadmill when some parts of the brain stem are stimulated. It has also been shown
that even in the complete absence of stimulus neural circuits exist in the spinal cord which can
produce oscillatory motor outputs. Similar results have been found in humans. In experiments
with newborn infants it has been shown that they are capable of producing stepping
movements when they are held upright. Also, in experiments with complete adult paraplegic
patients, it is shown that they are able to produce stepping movements when supported on a
moving treadmill. These results suggest that the same neural circuits which are capable of
autonomous rhythm generation invertebrates may be involved in human bipedal locomotion.

DEFINITION OF LOCOMOTION

Locomotion is defined as a traslatory progression of the body as a whole, produced by

coordinated, rotatory movements of the body segments as a result of combined and
coordinated actions of neuro, muscular and skeletal systems.
 ROLE OF DIFFERENT PARTS OF NERVOUS SYSTEM IN
HUMAN LOCOMOTION

Neural convergence

Convergence refers to multiple afferent inputs onto a single neuron or nucleus.

Afferent input refers to signals providing incoming information to the target neuron (includes
descending as well as peripheral sensory input).convergence occurs at every level of the
Central Nervous System. Segmentally, interneurons and motor neurons receive multiple
inputs. The complexity increases in supraspinal centers. Determining which input has greater
relative influence is a difficult task. The difficulty of the task is increased by the fact that the
relative influences of afferent inputs change within a situational context, the position of the
limbs, weight bearing status and anticipatory motor set. Convergence is an important concept
to bear in mind to understand about the inputs and outputs of various neural structures and
attempting to summarise function based on connectivity. Because of the complexity
introduced by the convergence, divergence and neuromodulation, however, descriptions of
neural connectivity alone may not be able accurately to determine what initiates, generates,
and maintains locomotion.

Peripheral receptors and afferents

The generation of locomotion depends on sensory input and the act of locomoting, in
turn, results in a wide array of sensory changes. Sensory afferent information derived from the
muscles can be considered among the most important. Control of bipedal locomotion requires
constant monitoring of muscle length and tension. This monitoring is provided by muscle
spindles, which are sensitive to changes in the muscle length and Golgi tendon organs (GTOs)
which are responsive to muscle tension. Spindles are innervated by gamma motor neurons.
The gamma motor neuron determines the sensitivity of the spindle to stretch. The sensitivity
of the spindle changes with the requirements of the movement. Dynamic changes in the
muscle length occur during the gait cycle. Therefore, the spindle not only serves as a feed
back receptor, but also as a feed-forward mechanism during human locomotion. GTOs are
contraction sensitive mechanoreceptors that are innervated by Ib afferent fibres. GTOs are
thought to alter the force out put of different muscles to constantly meet the requirements of
the ongoing movement. The timing of the locomotor rhythm is strongly influenced by group I
afferents.

Flexor reflex afferents

Flexor reflex afferents (FRAs) refers to a multisensorial and interneuronal reflex

system that appears to be at least partially responsible for the generation of locomotion. The
afferents include in this reflex pathway include mechanoreceptors, cutaneous afferents,
nociceptors, joint afferents and muscle afferents.
Activity in one motor neuronal pool generally results in inhibition of antagonist pools.
Alternating activity between various motor neuron pool forms the basis for the rhythmic
locomotor activity.
Brainstem locomotor regions

Electrical stimulation of the non human mammalian midbrain and brainstem causes
spontaneous locomotion. The three such centres are the mesencephalic locomotor region
(MLR), the pontine locomotor region (PLR), the subthalamic region (SLR). These centres
receive and process diverse descending and afferent input. Their efferent output is to spinal
CPGs.

The MLR receives afferent input from the basal ganglia, the limbic system, and the
sensorimotor cortex. It connects with the spinal circuitry via the reticulospinal tract. the MLR
appears to be an important relay station between the cortical limbic drives and the CPGs. By
the nature of the sympathetic inputs to the limbic systems, the MLR may be the centre by
which the “flight” component of the “fight or flight” sympathetic response becomes manifest.
The PLR is located caudal to the MLR and may actually be contiguous with it. some have
suggested that a pontomedullarylocomotor strip extends from the MLR to the upper cervical
spinal cord. Stimulation of the area changes postural tone and alters firing within the MLR.

If the SLR is disconnected from the cortical centres in cat, the animal can ambulate
spontaneously but loses all ability to avoid obstacles. The SLR, therefore, appears to be
necessary for the modulation of the locomotor pattern. A lesion immediately caudal to the
SLR in cats will result in the disappearance of all locomotion.

Cerebellum

The cerebellum is known to be involved in the control of quick limb movements

(Ghezand Thach, 2000). For example, patients with cerebellar damage are strongly impaired
in the control of multi-joint movements such as arm reaching and locomotion, although it is
possible for them to execute simple, single-joint movements. It has also been demonstrated in
experimental animals that the adaptation of quick eye movements is dependent on the
cerebellum. Accordingly, it has been hypothesized that the major role of the cerebellum is
both the temporal and spatial coordination of movements. The cerebellum receives
somatotopically organized input from the cerebral cortex (motor areas 4 and 6 visual and
auditory cortices) brainstem nuclei such as vestibular nuclei and the midbrain. The ventral and
dorsal spinocerebellartracts relay information from the peripheral muscle spindles, GTOs, and
joint afferents to the cerebellum during movement. Both tracts are physically active during
locomotion. The dorsal tracts appear to convey the information about the activity of
individual muscles. The ventral tract receives more diffuse input and may be involved in
comparing the descending copy of the motor program for the locomotion with the resultant
changes in the periphery.

Basal ganglia

The functions of basal ganglia with regard to movement and locomotion remain a mystery.
The basal ganglia are thought to process afferent information from the periphery and the
cerebral cortex and somehow impact motor planning. They play an important role in the
initiation and termination of movement. Some basal ganglia neuron fire before movement,
whereas other fire after movement and after neurons within the sensorymotor cortex have
already finished firing. Neurons within the basal ganglia respond to sensory inputs related to
movements.

Cerebral cortex

Known functions of the cerebral cortex with regard to locomotion include cognitive
aspects of the motor control, visuomotor coordination, and motor planning. Cortical neurons
are often activated before movement onset, typically fire phasically during locomotion. The
possible role of the cerebral cortex in the generation and coordination of human locomotion
remains controversial. The bipedal, plantigrade gait is uniquely a human activity. In addition
to increased equilibrium demands, bipedal gait involves increased anticipatory muscle
activations, the functional stretch reflex, and bodily responses to the unweighting of a limb
and single leg stance. All of these functions rely on cerebral centres.

THEORIES OF MOTOR CONTROL

Reflex Theory:

A reflex theory of motor control was first proposed in 1906 by Sir Charles
Sherrington, a neurophysiologist. This theory views movement as a combination or sequence
of reflexes. Complex movements were described in terms of compound reflexes and their
successive chaining. A stimulus is required at a receptor, and is conducted via a neural
pathway to an effector (muscle) which produces a motor response. This structure is known as
the reflex arc, and through feedback may produce the next stimulus for the next response
(chaining). Over the years, therapists have referred to this concept as a sensory-motor, cause-
effect system.

Hierarchical Theory:

Hierarchical theory was proposed in the 1920's and 1930's by several researchers who
used the continuing research of reflexes to make observations and interpretations regarding
the role of the higher brain centers as a controlling mechanism. Reflex and hierarchical
theories were combined into one, and are referred to as a reflex/hierarchical theory of motor
control. This theory views movement as emerging from reflex patterns that are controlled by
hierarchically organized levels of the central nervous system. This model uses a top-down
structure, in which higher centers control or inhibit activity of the lower centers. In the 1940's,
Gesell and McGraw used the reflex/hierarchical theory to describe infant maturation and
childhood development.

Ecological theory:

Ecological theory was first proposed by James Gibson in 1966, and expanded upon by
his students. This model is now known as the ecological approach to motor control. This
theory holds that all movements and actions are influenced or constrained by the environment.
Environmental information is necessary to shape or modify the characteristics of movement to
achieve specific actions or tasks. Whereas previous approaches viewed the individual as a
sensory-motor system, this new theory holds that it is not mere sensation that stimulates the
response. Of primary importance is the perception of the environmental factors and sensory
information that guide the individual to coordinate movements to accomplish a desired goal-
directed task. The individual must be viewed as organizing actions that are specific to the
desired task within the environment in which the task is being performed.

CENTRAL PATTERN GENERATORS (CPGs)

Central pattern generators are neuronal circuits that when activated can produce
rhythmic motor patterns such as walking, breathing, flying, and swimming in the absence of
sensory or descending inputs that carry specific timing information. General principles of the
organization of these circuits and their control by higher brain centers have come from the
study of smaller circuits found in invertebrates. Recent work on vertebrates highlights the
importance of neuro-modulatory control pathways in enabling spinal cord and brain stem
circuits to generate meaningful motor patterns. Because rhythmic motor patterns are easily
quantified and studied, central pattern generators will provide important testing grounds for
understanding the effects of numerous genetic mutations on behavior. Moreover, further
understanding of the modulation of spinal cord circuitry used in rhythmic behaviors should
facilitate the development of new treatments to enhance recovery after spinal cord damage.

FIG. 1. Schematic drawing of the organization of the principal sensory-central interactions in

locomotor pattern generation. A central pattern generating network (CPG) depicted in a
square box with 2 neurons A and B, interacting with each other, generates a basic rhythmic
output that drives sets of antagonistic motoneurons pools and muscles (combined in square
boxes and marked with letters in italics). Feedback about the resulting limb movement is
provided by sense organs. Sensory feedback can contribute either to the control of magnitude
of motor output or to the control of timing of motor activity.

STUDIES ON ANIMALS AND HUMAN BEINGS

Evidence for a spinal central pattern generator in humans.

Dimitrijevic MR, Gerasimenko Y, Pinter MM.

Department of Physical Medicine and Rehabilitation, Baylor College of Medicine, Houston,

Texas 77030, USA. agoshie@bcm.tmc.edu

Non-patterned electrical stimulation of the posterior structures of the lumbar spinal cord in
subjects with complete, long-standing spinal cord injury can induce patterned, locomotor-like
activity. We show that epidural spinal cord stimulation can elicit step-like EMG activity and
locomotor synergies in paraplegic subjects. An electrical train of stimuli applied over the
second lumbar segment with a frequency of 25 to 60 Hz and amplitude of 5-9 V was effective
in inducing rhythmic, alternating stance and swing phases of the lower limbs. This finding
suggests that spinal circuitry in humans has the capability of generating locomotor-like
activity even when isolated from brain control, and that externally controlled sustained
electrical stimulation of the spinal cord can replace the tonic drive generated by the brain.

(pmid: 9928325 [pubmed - indexed for medline])

SUMMARY

• By understanding neural control of skilled human locomotor behavior can serve to

guide further studies on locomotion by highlighting what is known and unknown
• It can be useful in interpreting deficits in locomotor behavior observed in patients and
evaluating rehabilative strategies
 • Thus vestibular system in postural control, visual system for locomotor system for
locomotor adaptation, neural and muscular adaptations to muscular fatigue during
locomotion and process of neural development that impact on the acquisition of
bipedal motion

CONCLUSION

Locomotion is essential for survival; it allows animals to acquire food, evade

predators, or find a mate.Many situations such as to escape from predators, crossing shallow
rivers quite possibly given rise to bipedilism Controlling locomotion is not an easy task.
Neural control of human locomotion is not yet fully understood. There is strong evidence
which suggests that isolated neural circuits called central pattern generators exist invertebrates
to produce rhythmic oscillatory patterns, which coupled with the dynamics of the body
produce locomotion. Experiments with cats have shown that a decerebrated cat can walk on a
treadmill when some parts of the brain stem are stimulated. It has also been shown that even
in the complete absence of stimulus neural circuits exist in the spinal cord which can produce
oscillatory motor outputs. Similar results have been found in humans. The existence of CPGs
is not clearly proved till date.