The second
way the spinal cord promotes homeostasis is by serving as an integrating center
for some reflexes. A reflex is a fast, automatic, unplanned sequence of actions
that occurs in response to a particular stimulus. Some reflexes are inborn, such
as pulling your hand away from a hot surface before you even feel that it is
hot. Other reflexes are learned or acquired. For instance, you learn many
reflexes while acquiring driving expertise. Slamming on the brakes in an
emergency is one example. When integration takes place in the spinal cord gray
matter, the reflex is a spinal reflex. An example is the familiar patellar reflex
(knee jerk). If integration occurs in the brain stem rather than the spinal
cord, the reflex is called a cranial reflex. An example is the tracking movements
of your eyes as you read this sentence. You are probably most aware of somatic
reflexes, which involve contraction of skeletal muscles. Equally important,
however, are the autonomic (visceral) reflexes, which generally are not
consciously perceived. They involve responses of smooth muscle, cardiac muscle,
and glands. The body functions such as heart rate,
digestion, urination, and defecation are controlled by the autonomic nervous
system through autonomic reflexes.
Nerve
impulses propagating into, through, and out of the CNS follow specific pathways,
depending on the kind of information, its origin, and its destination. The
pathway followed by nerve impulses that produce a reflex is a reflex arc (reflex
circuit). A reflex arc includes the following five functional components.
- Sensory receptor. The distal end of a sensory neuron (dendrite) or an associated sensory structure serves as a sensory receptor. It responds to a specific stimulus a change in the internal or external environment by producing a graded potential called a generator (or receptor) potential. If a generator potential reaches the threshold level of depolarization, it will trigger one or more nerve impulses in the sensory neuron.
- Sensory neuron. The nerve impulses propagate from the sensory receptor along the axon of the sensory neuron to the axon terminals, which are located in the gray matter of the spinal cord or brain stem.
- Integrating center. One or more regions of gray matter within the CNS act as an integrating center. In the simplest type of reflex, the integrating center is a single synapse between a sensory neuron and a motor neuron. A reflex pathway having only one synapse in the CNS is termed a monosynaptic reflex arc (mono- one). More often, the integrating center consists of one or more interneurons, which may relay impulses to other interneurons as well as to a motor neuron. A polysynaptic reflex arc (poly- many) involves more than two types of neurons and more than one CNS synapse.
COMPONENTS OF REFLEX CIRCUIT
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- Motor neuron. Impulses triggered by the integrating center propagate out of the CNS along a motor neuron to the part of the body that will respond.
- Effector. The part of the body that responds to the motor nerve impulse, such as a muscle or gland, is the effector. Its action is called a reflex. If the effector is skeletal muscle, the reflex is a somatic reflex. If the effector is smooth muscle, cardiac muscle, or a gland, the reflex is an autonomic (visceral) reflex.
Because reflexes are normally so predictable, they provide useful
information about the health of the nervous system and can greatly aid
diagnosis of disease. Damage or disease anywhere along its reflex arc can cause
a reflex to be absent or abnormal. For example, tapping the patellar ligament
normally causes reflex extension of the knee joint. Absence of the patellar
reflex could indicate damage of the sensory or motor neurons, or a spinal cord
injury in the lumbar region. Somatic reflexes generally can be tested simply by
tapping or stroking the body surface. Next, we examine four important somatic
spinal reflexes: the stretch reflex, the tendon reflex, the flexor (withdrawal)
reflex, and the crossed extensor reflex.
The Stretch Reflex
A stretch reflex causes contraction of a skeletal muscle (the effector) in
response to stretching of the muscle. This type of reflex occurs via a
monosynaptic reflex arc. The reflex can occur by activation of a single sensory
neuron that forms one synapse in the CNS with a single motor neuron. Stretch
reflexes can be elicited by tapping on tendons attached to muscles at the elbow,
wrist, knee, and ankle joints. An example of a stretch reflex is the patellar
reflex (knee jerk); a stretch reflex operates as follows:
1. Slight stretching of a muscle
stimulates sensory receptors in the muscle called muscle spindles. The spindles
monitor changes in the length of the muscle.
2. In response to being stretched, a
muscle spindle generates one or more nerve impulses that propagate along a
somatic sensory neuron through the posterior root of the spinal nerve and into
the spinal cord.
3. In the spinal cord (integrating
center), the sensory neuron makes an excitatory synapse with and thereby
activates a motor neuron in the anterior gray horn.
4. If the excitation is strong enough,
one or more nerve impulses arise in the motor neuron and propagate along its
axon, which extends from the spinal cord into the anterior root and through
peripheral nerves to the stimulated muscle. The axon terminals of the motor
neuron form neuromuscular junctions (NMJs) with skeletal muscle fibers of the
stretched muscle.
5. Acetylcholine released by nerve
impulses at the NMJs triggers one or more muscle action potentials in the
stretched muscle (effector), and the muscle contracts. Thus, muscle stretch is
followed by muscle contraction, which relieves the stretching.
COMPONENTS OD STRETCH REFLEX
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In the reflex
arc just described, sensory nerve impulses enter the spinal cord on the same
side from which motor nerve impulses leave it. This arrangement is called an
ipsilateral reflex ( same side). All monosynaptic reflexes are ipsilateral. In
addition to the large-diameter motor neurons that innervate typical skeletal
muscle fibers, smaller-diameter motor neurons innervate smaller, specialized
muscle fibers within the muscle spindles themselves. The brain regulates muscle
spindle sensitivity through pathways to these smaller motor neurons. This
regulation ensures proper muscle spindle signaling over a wide range of muscle
lengths during voluntary and reflex contractions. By adjusting how vigorously a
muscle spindle responds to stretching, the brain sets an overall level of
muscle tone, which is the small degree of contraction present while the muscle
is at rest. Because the stimulus for the stretch reflex is stretching of muscle,
this reflex helps avert injury by preventing overstretching of muscles. Although
the stretch reflex pathway itself is monosynaptic (just two neurons and one
synapse), a polysynaptic reflex arc to the antagonistic muscles operates at the
same time. This arc involves three neurons and two synapses. Axon collateral
(branch) from the muscle spindle sensory neuron also synapses with an
inhibitory interneuron in the integrating center. In turn, the interneuron
synapses with and inhibits a motor neuron that normally excites the antagonistic
muscles. Thus, when the stretched muscle contracts during a stretch reflex,
antagonistic muscles that oppose the contraction relax. This type of
arrangement, in which the components of a neural circuit simultaneously cause
contraction of one muscle and relaxation of its antagonists, is termed
reciprocal innervation. Reciprocal innervation prevents conflict between
opposing muscles and is vital in coordinating body movements. Axon collaterals
of the muscle spindle sensory neuron also relay nerve impulses to the brain
over specific ascending pathways. In this way, the brain receives input about
the state of stretch or contraction of skeletal muscles, enabling it to
coordinate muscular movements. The nerve impulses that pass to the brain also
allow conscious awareness that the reflex has occurred.
The stretch
reflex can also help maintain posture. For example, if a standing person begins
to lean forward, the gastrocnemius and other calf muscles are stretched.
Consequently, stretch reflexes are initiated in these muscles, which cause them
to contract and reestablish the body’s upright posture. Similar types of
stretch reflexes occur in the muscles of the shin when a standing person begins
to lean backward.
The Tendon Reflex
The stretch
reflex operates as a feedback mechanism to control muscle length by causing
muscle contraction. In contrast, the tendon reflex operates as a feedback
mechanism to control muscle tension by causing muscle relaxation before muscle force
becomes so great that tendons might be torn. Although the tendon reflex is less
sensitive than the stretch reflex, it can override the stretch reflex when
tension is great, making you drop a very heavy weight, for example. Like the
stretch reflex, the tendon reflex is ipsilateral. The sensory receptors for this
reflex are called tendon (Golgi tendon) organs, which lie within a tendon near
its junction with a muscle. In contrast to muscle spindles, which are sensitive
to changes in muscle length, tendon organs detect and respond to changes in
muscle tension that are caused by passive stretch or muscular contraction.
A tendon
reflex operates as follows:
1. As the
tension applied to a tendon increases, the tendon organ (sensory receptor) is
stimulated (depolarized to threshold).
2. Nerve impulses arise and propagate
into the spinal cord along a sensory neuron.
3. Within the spinal cord
(integrating center),the sensory neuron activates an inhibitory interneuron
that synapses with a motor neuron.
4. The inhibitory neurotransmitter inhibits
(hyperpolarizes) the motor neuron, which then generates fewer nerve impulses.
5. The muscle relaxes and relieves excess tension.
Thus, as
tension on the tendon organ increases, the frequency of inhibitory impulses
increases; inhibition of the motor neurons to the muscle developing excess
tension (effector) causes relaxation of the muscle. In this way, the tendon
reflex protects the tendon and muscle from damage due to excessive tension. Note that the sensory neuron from the tendon organ also synapses
with an excitatory interneuron in the spinal cord. The excitatory interneuron
in turn synapses with motor neurons controlling antagonistic muscles. Thus,
while the tendon reflex brings about relaxation of the muscle attached to the
tendon organ, it also triggers contraction of antagonists. Here we have another
example of reciprocal innervation. The sensory neuron also relays nerve
impulses to the brain by way of sensory tracts, thus informing the brain about
the state of muscle tension throughout the body.
The Flexor
and Crossed Extensor Reflexes
Another reflex involving a polysynaptic reflex arc
results when, for instance, you step on a tack. In response to such a painful
stimulus, you immediately withdraw your leg. This reflex, called the flexor or
withdrawal reflex, operates as follow:
- Stepping on a tack stimulates the dendrites (sensory receptor) of a pain-sensitive neuron.
- This sensory neuron then generates nerve impulses, which propagate into the spinal cord.
- Within the spinal cord (integrating center),the sensory neuron activates interneurons that extend to several spinal cord segments.
- The interneurons activate motor neurons in several spinal cord segments. As a result, the motor neurons generate nerve impulses, which propagate toward the axon terminals.
- Acetylcholine released by the motor neurons causes the flexor muscles in the thigh (effectors) to contract, producing withdrawal of the leg. This reflex is protective because contraction of flexor muscles moves a limb away from the source of a possibly damaging stimulus.
FLEXOR (WITHDRAWAL) REFLEX
The flexor
reflex, like the stretch reflex, is ipsilateral—the incoming and outgoing
impulses propagate into and out of the same side of the spinal cord. The flexor
reflex also illustrates another feature of polysynaptic reflex arcs. Moving your
entire lower or upper limb away from a painful stimulus involves contraction of
more than one muscle group. Hence, several motor neurons must simultaneously
convey impulses to several limb muscles. Because nerve impulses from one
sensory neuron ascend and descend in the spinal cord and activate interneurons
in several segments of the spinal cord, this type of reflex is called an
intersegmental reflex arc (inter- between). Through intersegmental reflex arcs, a
single sensory neuron can activate several motor neurons, thereby stimulating
more than one effector. The monosynaptic stretch reflex, in contrast, involves
muscles receiving nerve impulses from one spinal cord segment only.
COMPONENTS OF FLEXOR REFLEX |
Something
else may happen when you step on a tack: You may start to lose your balance as
your body weight shifts to the other foot. Besides initiating the flexor reflex
that causes you to withdraw the limb, the pain impulses from stepping on the
tack also initiate a crossed extensor reflex to help you maintain your balance;
it operates as follows:
- Stepping on a tack stimulates the sensory receptor of a painsensitive neuron in the right foot.
- This sensory neuron then generates nerve impulses, which propagate into the spinal cord.
- Within the spinal cord (integrating center),the sensory neuron activates several interneurons that synapse with motor neurons on the left side of the spinal cord in several spinal cord segments. Thus, incoming pain signals cross to the opposite side through interneurons at that level, and at several levels above and below the point of entry into the spinal cord.
- The interneurons excite motor neurons in several spinal cord segments that innervate extensor muscles. The motor neurons in turn generate more nerve impulses, which propagate toward the axon terminals.
- Acetylcholine released by the motor neurons causes extensor muscles in the thigh (effectors) of the unstimulated left limb to contract, producing extension of the left leg. In this way, weight can be placed on the foot that must now support the entire body. A comparable reflex occurs with painful stimulation of the left lower limb or either upper limb.
Unlike the
flexor reflex, which is an ipsilateral reflex, the crossed extensor reflex involves
a contralateral reflex arc Sensory impulses enter one side of the spinal cord
and motor impulses exit on the opposite side. Thus, a crossed extensor reflex
synchronizes the extension of the contralateral limb with the withdrawal
(flexion) of the stimulated limb. Reciprocal innervation also occurs in both the
flexor reflex and the crossed extensor reflex. In the flexor reflex, when the flexor
muscles of a painfully stimulated lower limb are contracting, the extensor
muscles of the same limb are relaxing to some degree. If both sets of muscles
contracted at the same time, the two sets of muscles would pull on the bones in
opposite directions, which might immobilize the limb. Because of reciprocal
innervation, one set of muscles contracts while the other relaxes.
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