THE SPINAL CORD & SPINAL NERVE

Tutik ermawati

Gross anatomi of spinal cord

The adult spinal cord measures approximately 45 cm (18 in.) in length and has a maximum width of roughly 14 mm (0.55 in.). The posterior (dorsal) surface of the spinal cord bears a shallow longitudinal groove, the posterior median sulcus. The anterior median fissure is a deeper groove along the anterior (ventral) surface. The amount of gray matter is greatest in segments of the spinal cord that deal with the sensory and motor control of the limbs. These areas are expanded, forming the enlargements of the spinal cord. The cervical enlargement supplies nerves to the shoulder girdles and upper limbs; the lumbar enlargement provides innervation to structures of the pelvis and lower limbs. Inferior to the lumbar enlargement, the spinal cord becomes tapered and conical; this region is the conus medullaris. The filum terminale ("terminal thread"), a slender strand of fibrous tissue, extends from the inferior tip of the conus medullaris. It continues along the length of the vertebral canal as far as the second sacral vertebra. There it provides longitudinal support to the spinal cord as a component of the coccygeal ligament.

The entire spinal cord can be divided into 31 segments on the basis of the origins of the spinal nerves. Each segment is identified by a letter and number designation, the same method used to identify vertebrae. Every spinal segment is associated with a pair of dorsal root ganglia. These ganglia, situated near the spinal cord, contain the cell bodies of sensory neurons. The axons of these neurons form the dorsal roots, which bring sensory information into the spinal cord. A pair of ventral roots contains the axons of motor neurons that extend into the periphery to control somatic and visceral effectors. On both sides, the dorsal and ventral roots of each segment pass between the vertebral canal and the periphery at the intervertebral foramen between successive vertebrae. The dorsal root ganglion lies between the pedicles of the adjacent vertebrae.

Distal to each dorsal root ganglion, the sensory and motor roots are bound together into a single spinal nerve. Spinal nerves are classified as mixed nervesthat is, they contain both afferent (sensory) and efferent (motor) fibers. There are 31 pairs of spinal nerves, each identified by its association with adjacent vertebrae. For example, we may speak of "cervical spinal nerves" or even "cervical nerves" when we make a general reference to spinal nerves of the neck. When we indicate specific spinal nerves, it is customary to give them a regional number, as indicated in Figure 13-1 . Each spinal nerve inferior to the first thoracic vertebra takes its name from the vertebra immediately preceding it. Thus, spinal nerve T1 emerges immediately inferior to vertebra T1 , spinal nerve T2 follows vertebra T2 , and so forth. The arrangement differs in the cervical region, because the first pair of spinal nerves, C1 , passes between the skull and the first cervical vertebra. For this reason, each cervical nerve takes its name from the vertebra immediately following it. In other words, cervical nerve C2 precedes vertebra C2 , and the same system is used for the rest of the cervical series. The transition from one numbering system to another occurs between the last cervical vertebra and first thoracic vertebra. Because the spinal nerve lying between them has been designated C8 , there are only seven cervical vertebrae but eight cervical nerves.

The spinal cord continues to enlarge and elongate until an individual is approximately 4 years old. Up to that time, enlargement of the spinal cord keeps pace with the growth of the vertebral column. Throughout this period, the ventral and dorsal roots are very short, and they enter the intervertebral foramina immediately adjacent to their spinal segment. After age 4, the vertebral column continues to elongate but the spinal cord does not. This vertebral growth carries the dorsal roots and spinal nerves farther and farther from their original positions relative to the spinal cord. As a result, the dorsal and ventral roots gradually elongate, and the correspondence between spinal segment and the vertebral segment is lost. For example, in adults the sacral segments of the spinal cord are at the level of vertebrae L1L2. Because the adult spinal cord extends only to the level of the first or second lumbar vertebra, the dorsal and ventral roots of spinal segments L2 to S5 extend inferiorly, past the inferior tip of the conus medullaris. When seen in gross dissection, the filum terminale and the long ventral and dorsal roots resemble a horse's tail. As a result, early anatomists called this complex the cauda equina.

SPINAL MENINGES

The vertebral column and its surrounding ligaments, tendons, and muscles isolate the spinal cord from the external environment. The delicate neural tissues must also be protected from damaging contacts with the surrounding bony walls of the vertebral canal. The spinal meninges, a series of specialized membranes surrounding the spinal cord, provide the necessary physical stability and shock absorption. Blood vessels branching within these layers deliver oxygen and nutrients to the spinal cord. The spinal meninges consist of three layers: (1) the dura mater, (2) the arachnoid, (3) the pia mater.

At the foramen magnum of the skull, the spinal meninges are continuous with the cranial meninges, which surround the brain. Bacterial or viral infection can cause meningitis, or inflammation of the meningeal membranes. Meningitis is dangerous because it can disrupt the normal circulatory and cerebrospinal fluid supplies, damaging or killing neurons and neuroglia in the affected areas. Although an initial diagnosis may specify the meninges of the spinal cord (spinal meningitis) or brain (cerebral meningitis), in later stages the entire meningeal system is usually affected.

The Dura Mater

The tough, fibrous dura mater is the layer that forms the outermost covering of the spinal cord. This layer contains dense collagen fibers that are oriented along the longitudinal axis of the spinal cord. Between the dura mater and the walls of the vertebral canal lies the epidural space, a region that contains loose connective tissue, blood vessels, and a protective padding of adipose tissue. The spinal dura mater does not have extensive, firm connections to the surrounding vertebrae. Longitudinal stability is provided by localized attachment sites at either end of the vertebral canal. Cranially, the outer layer of the spinal dura mater fuses with the periosteum of the occipital bone around the margins of the foramen magnum. There the spinal dura mater becomes continuous with the cranial dura mater. Within the sacral canal, the spinal dura mater tapers from a sheath to a dense cord of collagen fibers that blend with components of the filum terminale to form the coccygeal ligament

The coccygeal ligament continues along the sacral canal, ultimately blending into the periosteum of the coccyx. Lateral support for the spinal dura mater is provided by loose connective tissue and adipose tissue within the epidural space. In addition, this dura mater extends between adjacent vertebrae at each intervertebral foramen, fusing with the connective tissues that surround the spinal nerves. Injecting an anesthetic into the epidural space of the spinal cord will affect only the spinal nerves in the immediate area of the injection. The result is an epidural blocka temporary sensory loss or a sensory and motor paralysis, depending on the anesthetic selected. Epidural blocks in the inferior lumbar or sacral regions may be used to control pain during childbirth. Spinal Anesthesia

The Arachnoid

In most anatomical and histological preparations, a narrow subdural space separates the dura mater from deeper meningeal layers. It is likely, however, that in life no such space exists and that the inner surface of the dura mater is in contact with the outer surface of the arachnoid, the middle meningeal layer. The inner surface of the dura mater and the outer surface of the arachnoid are covered by simple squamous epithelia. The arachnoid includes this epithelium, called the arachnoid membrane, and the arachnoid trabeculae, a delicate network of collagen and elastic fibers that extends between the arachnoid membrane and the outer surface of the pia mater.

The intervening region is called the subarachnoid space. It is filled with cerebrospinal fluid (CSF), which acts as a shock absorber and a diffusion medium for dissolved gases, nutrients, chemical messengers, and waste products. The spinal arachnoid extends inferiorly as far as the filum terminale, and the dorsal and ventral roots of the cauda equina lie within the fluid-filled subarachnoid space. In adults, the withdrawal of cerebrospinal fluid, a procedure known as a spinal tap, involves the insertion of a needle into the subarachnoid space in the inferior lumbar region. (See the clinical discussion "Spinal Taps and Myelography.") This placement avoids damage to the spinal cord. Spinal taps are performed when CNS infection is suspected or to diagnose severe back pain, headaches, disc problems, and some types of strokes.

The Pia Mater

The subarachnoid space bridges the gap between the arachnoid epithelium and the innermost meningeal layer, the pia mater. The pia mater consists of a meshwork of elastic and collagen fibers that is firmly bound to the underlying neural tissue. These connective tissue fibers are extensively interwoven with those that span the subarachnoid space, firmly binding the arachnoid to the pia mater. The blood vessels servicing the spinal cord run along the surface of the spinal pia mater, within the subarachnoid space. Along the length of the spinal cord, paired denticulate ligaments extend from the pia mater through the arachnoid to the dura mater. Denticulate ligaments, which originate along either side of the spinal cord, prevent lateral (side-toside) movement. The dural connections at the foramen magnum and the coccygeal ligament prevent longitudinal (superior/inferior) movement. The spinal meninges accompany the dorsal and ventral roots as these roots pass through the intervertebral foramina. the meningeal membranes are continuous with the connective tissues that surround the spinal nerves and their peripheral branches

SECTIONAL ANATOMY OF THE SPINAL CORD

To understand the functional organization of the spinal cord, you must become familiar with its sectional organization. The anterior median fissure and the posterior median sulcus mark the division between left and right sides of the spinal cord. The superficial white matter contains large numbers of myelinated and unmyelinated axons. The gray matter, dominated by the cell bodies of neurons, neuroglia, and unmyelinated axons, surrounds the narrow central canal and forms an H or butterfly shape. The projections of gray matter toward the outer surface of the spinal cord are called horns.

Organization of Gray Matter

The cell bodies of neurons in the gray matter of the spinal cord are organized into functional groups called nuclei. Sensory nuclei receive and relay sensory information from peripheral receptors. Motor nuclei issue motor commands to peripheral effectors. Although sensory and motor nuclei appear rather small in transverse section, they may extend for a considerable distance along the length of the spinal cord. A frontal section along the length of the central canal of the spinal cord separates the sensory (posterior, or dorsal) nuclei from the motor (anterior, or ventral) nuclei. The posterior gray horns contain somatic and visceral sensory nuclei, whereas the anterior gray horns contain somatic motor nuclei. The lateral gray horns, located only in the thoracic and lumbar segments, contain visceral motor nuclei. The gray commissures (commissura, a joining together) posterior to and anterior to the central canal contain axons that cross from one side of the cord to the other before they reach a destination in the gray matter.

shows the relationship between the function of a particular nucleus (sensory or motor) and its relative position in the gray matter of the spinal cord. The nuclei within each gray horn are also organized. For example, the anterior gray horns of the cervical enlargement contain nuclei whose motor neurons control the muscles of the upper limbs. On each side of the spinal cord, in medial to lateral sequence, are somatic motor nuclei that control (1) muscles that position the pectoral girdle (2) muscles that move the arm (3) muscles that move the forearm and hand (4) muscles that move the hand and fingers. Within each of these regions, the motor neurons that control flexor muscles are grouped separately from those that control extensor muscles. Because the spinal cord is so highly organized, we can predict the muscles that will be affected by damage to a specific area of gray matter.

Organization of White Matter

The white matter on each side of the spinal cord can be divided into three regions called columns, or funiculi. The posterior white columns lie between the posterior gray horns and the posterior median sulcus. The anterior white columns lie between the anterior gray horns and the anterior median fissure. The anterior white columns are interconnected by the anterior white commissure, a region where axons cross from one side of the spinal cord to the other. The white matter between the anterior and posterior columns on each side makes up the lateral white column. Each column contains tracts whose axons share functional and structural characteristics. A tract, or fasciculus, is a bundle of axons in the CNS that are relatively uniform with respect to diameter, myelination, and conduction speed. All the axons within a tract relay the same type of information (sensory or motor) in the same direction. Short tracts carry sensory or motor signals between segments of the spinal cord, and longer tracts connect the spinal cord with the brain. Ascending tracts carry sensory information toward the brain, and descending tracts convey motor commands to the spinal cord.

SPINAL NERVE

A series of connective tissue layers surrounds each spinal nerve and continues along all its peripheral branches. These layers, best seen in sectional view, are comparable to those associated with skeletal muscles. The epineurium, or outermost layer, consists of a dense network of collagen fibers. The fibers of the perineurium, the middle layer, extend inward from the epineurium. These connective tissue partitions divide the nerve into a series of compartments that contain bundles of axons, or fascicles. Delicate connective tissue fibers of the endoneurium, the innermost layer, extend from the perineurium and surround individual axons. Arteries and veins penetrate the epineurium and branch within the perineurium. Capillaries leaving the perineurium branch in the endoneurium and supply the axons and Schwann cells of the nerve and the fibroblasts of the connective tissues.

PERIPHERAL DISTRIBUTION OF SPINAL NERVES

The spinal nerve forms just lateral to the intervertebral foramen, where the dorsal and ventral roots unite. Let us follow the nerve's distribution in the periphery. In the thoracic and superior lumbar regions (segments T1L2 ), the first branch from the spinal nerve carries visceral motor fibers to a nearby autonomic ganglion. This ganglion is part of the sympathetic division of the autonomic nervous system (ANS). (The sympathetic division is, among its other functions, responsible for elevating the metabolic rate and for increasing alertness.) Because preganglionic axons are myelinated, this branch has a light color and is known as the white ramus ("branch"). Postganglionic fibers innervating glands and smooth muscles in the body wall or limbs return from the ganglia to rejoin the spinal nerve. These fibers are unmyelinated and have a darker color; they form the gray ramus. The gray ramus is typically proximal to the white ramus. The gray and white rami are collectively termed the rami communicantes, or "communicating branches." Postganglionic fibers innervating visceral organs in the thoracic cavity form a series of separate sympathetic nerves.

The dorsal ramus of each spinal nerve is a bundle of axons that provide sensory and motor innervation to the skin and muscles of the back. The axons in the relatively large ventral ramus supply the ventrolateral body surface, structures in the body wall, and the limbs. The specific region of the skin surface monitored by a single pair of spinal nerves is known as a dermatome. Each pair of spinal nerves services its own dermatome, although the precise boundaries of adjacent dermatomes overlap to some degree. Dermatomes are clinically important, because damage or infection of a spinal nerve or dorsal root ganglion will produce a characteristic loss of sensation in the skin. For example, in the condition shingles, a virus infects dorsal root ganglia, causing a painful rash whose distribution corresponds to that of the affected sensory nerves.

NERVE PLEXUSES

The simple distribution pattern of dorsal and ventral rami, applies to spinal nerves T2T12. But in segments controlling the skeletal musculature of the neck, upper limbs, or lower limbs, the situation is more complicated. During development, small skeletal muscles innervated by different ventral rami typically fuse to form larger muscles with compound origins. The anatomical distinctions between the component muscles may disappear, but separate ventral rami continue to provide sensory innervation and motor control to each part of the compound muscle. As they converge, the ventral rami of adjacent spinal nerves blend their fibers, producing a series of compound nerve trunks. Such a complex interwoven network of nerves is a nerve plexus There are four major plexuses: (1) the cervical plexus, (2) the brachial plexus, (3) the lumbar plexus, and (4) the sacral plexus.

The Cervical Plexus

The cervical plexus consists of the ventral rami of spinal nerves C1C5. Its branches innervate the muscles of the neck and extend into the thoracic cavity, where they control the diaphragmatic muscles. The phrenic nerve, the major nerve of the cervical plexus, provides the entire nerve supply to the diaphragm, a key respiratory muscle. Other branches of this nerve plexus are distributed to the skin of the neck and the upper part of the chest.

The Brachial Plexus

The brachial plexus innervates the pectoral girdle and upper limb, with contributions from the ventral rami of spinal nerves C5T1. The nerves that form this plexus originate from trunks and cords. Trunks are large bundles of axons contributed by several spinal nerves. Cords are smaller branches that originate at trunks. Both trunks and cords are named according to their location relative to the axillary artery, a large artery supplying the upper limb. Hence we have superior, middle, and inferior trunks and lateral, medial, and posterior cords. The lateral cord forms the musculocutaneous nerve exclusively and, together with the medial cord, contributes to the median nerve. The ulnar nerve is the other major nerve of the medial cord. The posterior cord gives rise to the axillary nerve and the radial nerve.

The Lumbar and Sacral Plexuses

The lumbar plexus and the sacral plexus arise from the lumbar and sacral segments of the spinal cord. The nerves arising at these plexuses innervate the pelvic girdle and lower limbs. The lumbar plexus contains axons from the ventral rami of spinal nerves T12 L4 . The major nerves of this plexus are the genitofemoral nerve, the lateral femoral cutaneous nerve, and the femoral nerve. The sacral plexus contains axons from the ventral rami of spinal nerves L4S4 . Two major nerves arise at this plexus: the sciatic nerve and the pudendal nerve. The sciatic nerve passes posterior to the femur, deep to the long head of the biceps femoris muscle. As it approaches the knee, the sciatic nerve divides into two branches--the peroneal nerve and the tibial nerve. The sural nerve, formed by branches of the peroneal nerve, is a sensory nerve innervating the lateral portion of the foot. A section of this nerve is often removed for use in nerve grafts.

REFLEXES

Conditions inside or outside the body can change rapidly and unexpectedly. Reflexes are rapid, automatic responses to specific stimuli. Reflexes preserve homeostasis by making rapid adjustments in the function of organs or organ systems. The response shows little variability; each time a particular reflex is activated, it usually produces the same motor response. Here we consider neural reflexes, in which sensory fibers deliver information to the CNS and motor fibers carry motor commands to peripheral effectors.

THE REFLEX ARC

The "wiring" of a single reflex is called a reflex arc. A reflex arc begins at a receptor and ends at a peripheral effector, such as a muscle fiber or a gland cell. the five steps in a neural reflex: step 1: The Arrival of a Stimulus and Activation of a Receptor. A receptor is a specialized cell or the dendrites of a sensory neuron. Receptors are sensitive to physical or chemical changes in the body or to the external environment. When you lean on a tack, pain receptors in the palm of your hand are activated. Pain receptors are the dendrites of sensory neurons; they respond to stimuli that cause or accompany tissue damage.

step 2: The Activation of a Sensory Neuron. The stimulation of pain receptors leads to the formation and propagation of action potentials along the axons of sensory neurons. This information reaches your spinal cord by way of a dorsal root. In our example, steps 1 and 2 involve the same cell. However, the two steps may involve different cells. For example, reflexes triggered by loud sounds begin when receptor cells in the inner ear release neurotransmitters that stimulate sensory neurons.

step 3: Information Processing. Information processing begins when a neurotransmitter released by synaptic knobs of each sensory neuron affects the postsynaptic membrane of an interneuron, producing an excitatory postsynaptic potential (EPSP). This stimulus is integrated with other stimuli arriving at the post-synaptic neuron at that moment. In a neuronal pool, the next step may coordinate an appropriate motor response by stimulating some motor neurons and inhibiting others. Interneurons are not always involved in a reflex arc; the sensory neuron may innervate a motor neuron directly. In that case, the motor neuron performs the information processing as summation occurs at the axon hillock.

step 4: The Activation of a Motor Neuron. The axons of the stimulated motor neurons carry action potentials into the periphery--in this example, through the ventral root of a spinal nerve. step 5: The Response of a Peripheral Effector. The release of neurotransmitter by the motor neurons at synaptic knobs then leads to a response by a peripheral effector in this case, a skeletal muscle whose contraction pulls your hand away from the tack.

A reflex response generally removes or opposes the original stimulus; here, the contracting muscle pulls your hand away from the painful stimulus. This reflex arc is therefore an example of negative feedback. By opposing potentially harmful changes in the internal or external environment, reflexes play an important role in homeostatic maintenance. The immediate reflex response is typically not the only response to a stimulus. In this example, you might wince, say "Ouch," and shake your hand. These other responses, which are directed by your brain, involve multiple synapses and take longer to organize and coordinate.

CLASSIFICATION OF REFLEXES

Reflexes are classified by (1) their development (2) the site of information processing (3) the nature of the resulting motor response (4) the complexity of the neural circuit involved. These categories are not mutually exclusive. They represent different ways of describing a single reflex

Development of Reflexes

Innate reflexes result from the connections that form between neurons during development. Such reflexes generally appear in a predictable sequence, from the simplest reflex responses (withdrawal from pain) to more-complex motor patterns (chewing, suckling, or tracking objects with the eyes). The neural connections responsible for basic motor patterns of an innate reflex are genetically or developmentally programmed. Examples include the reflexive removal of your hand from a hot stovetop and blinking when your eyelashes are touched. More-complex, learned motor patterns are called acquired reflexes. An experienced driver steps on the brake when trouble appears ahead; a professional skier makes rapid, automatic adjustments in body position while racing. These motor responses are rapid and automatic, but they were learned rather than preestablished. Such reflexes are enhanced by repetition. The distinction between innate and acquired reflexes is not absolute. Some people can learn motor patterns more quickly than others can, and the differences probably have a genetic basis.

Most reflexes, whether innate or acquired, can be modified over time or suppressed through conscious effort. For example, while walking a tightrope over the Grand Canyon, you might ignore a bee sting on your hand, although under other circumstances you would probably withdraw your hand immediately, with shouting and thrashing.

Processing Sites

In a spinal reflex, the important interconnections and processing events occur in the spinal cord. We will discuss these reflexes in detail in the next section. Reflexes processed in the brain are called cranial reflexes. For example, the reflex movements in response to a sudden loud noise or bright light are cranial reflexes directed by nuclei in the brain.

Nature of the Response

Somatic reflexes provide a mechanism for the involuntary control of the muscular system. Visceral reflexes, or autonomic reflexes, control the activities of other systems. The movements directed by somatic reflexes are not delicate or precise. You might therefore wonder why they exist at all, because we have voluntary control over the same muscles. Somatic reflexes are absolutely vital, primarily because they are immediate. Making decisions and coordinating voluntary responses take time, and in an emergency situationyou slip while descending a flight of stairs or lean your hand against a knife edgeany delay increases the likelihood of a severe injury. Thus, somatic reflexes provide a rapid response that, if necessary, can later be modified by voluntary motor commands.

Complexity of the Circuit

In the simplest reflex arc, a sensory neuron synapses directly on a motor neuron, which serves as the processing center. Such a reflex is a monosynaptic reflex. Transmission across a chemical synapse always involves a synaptic delay; but, with only one synapse, the delay between the stimulus and the response is minimized. Most other types of reflexes have at least one interneuron between the sensory neuron and the motor neuron. These polysynaptic reflexes have a longer delay between stimulus and response. The length of the delay is proportional to the number of synapses involved. Polysynaptic reflexes can produce far more complicated responses than monosynaptic reflexes can, because the interneurons can control several muscle groups.

SPINAL REFLEXES

Spinal reflexes range in complexity from simple monosynaptic reflexes involving a single segment of the spinal cord to polysynaptic reflexes that involve many segments. In the most complicated spinal reflexes, called intersegmental reflex arcs, many segments interact to produce a coordinated, highly variable motor response.

MONOSYNAPTIC REFLEXES

In a monosynaptic reflex, there is little delay between sensory input and motor output. These reflexes control the most-rapid, stereotyped motor responses of the nervous system to specific stimuli. The best-known monosynaptic reflex is the stretch reflex.

The Stretch Reflex The stretch reflex provides automatic regulation of skeletal muscle length. The stimulus (increasing muscle length) activates a sensory neuron, which triggers an immediate motor response (contraction of the stretched muscle) that counteracts the stimulus. Action potentials traveling toward and away from the spinal cord are conducted along large, myelinated, Type A fibers. The entire reflex is completed within 2040 msec.

The patellar reflex, or knee jerk reflex, is a familiar example of a stretch reflex. This reflex is triggered by stretching the quadriceps muscles, which extend the knee. You can demonstrate this reflex by tapping on the patellar tendon of a flexed knee when the thigh and leg muscles are relaxed. A tap on the patellar ligament stretches muscle spindles in the quadriceps group. This stretching produces a sudden burst of activity in the sensory neurons that monitor these muscle spindles, so muscle tone in the quadriceps group increases rapidly. The rise is so sudden that it usually produces a noticeable kick. As the muscles contract, the muscle spindles return to their resting length; the rate of action potential generation in the sensory neurons declines. This decline causes a drop in muscle tone, and the leg then swings back.

Muscle Spindles The sensory receptors involved in the stretch reflex are muscle spindles. Each muscle spindle consists of a bundle of small, specialized skeletal muscle fibers called intrafusal muscle fibers. The muscle spindle is surrounded by larger extrafusal muscle fibers, responsible for the resting muscle tone and, at greater levels of stimulation, for the contraction of the entire muscle. Each intrafusal fiber is innervated by both sensory and motor neurons. The dendrites of the sensory neuron spiral around the central portion of the intrafusal fiber. Axons from a separate population of spinal motor neurons form neuromuscular junctions on either end of this fiber. Motor neurons innervating intrafusal fibers are called gamma motor neurons; their axons are called gamma efferents. An intrafusal fiber has one set of myofibrils at each end. Instead of running the length of the fiber, each set of myofibrils is attached to the sarcolemma in the central region that is monitored by the sensory neuron.

The sensory neuron is always active, conducting impulses to the CNS. The axon enters the CNS in a dorsal root and synapses on motor neurons in the anterior gray horn of the spinal cord. Collaterals distribute the information to the brain, providing information about the state of the muscle spindle. Stretching the central portion of the intrafusal fiber distorts the dendrites and stimulates the sensory neuron, increasing the frequency of action potential generation. Compressing the central portion inhibits the sensory neuron, decreasing the frequency of action potential generation. The axon of the sensory neuron synapses on CNS motor neurons that control the extrafusal muscle fibers of the same muscle. An increase in stimulation of the sensory neuron, caused by stretching of the intrafusal fiber, will increase stimulation to the surrounding extrafusal fibers, so muscle tone increases. A decrease in the stimulation of the sensory neuron, due to compression of the intrafusal fiber, will lead to a decrease in the stimulation of extrafusal fibers and in turn to a drop in muscle tone.

When a skeletal muscle is stretched, muscle spindles elongate and muscle tone increases. This increase provides automatic resistance that reduces the chance of muscle damage due to overstretching. When a skeletal muscle is compressed, muscle spindles are also compressed, and muscle tone decreases. This decrease helps reduce resistance to the movement under way. Many stretch reflexes are postural reflexes, reflexes that maintain normal upright posture. For example, standing involves a cooperative effort of many muscle groups. Some of these muscles work in opposition to one another, exerting forces that keep the body's weight balanced over the feet. If the body leans forward, stretch receptors in the calf muscles are stimulated; those muscles respond by contracting. This contraction returns the body to an upright position. If the muscles overcompensate and the body begins to lean back, the calf muscles relax. But then stretch receptors in muscles of the shins and thighs are stimulated, and the problem is corrected immediately.

Postural muscles generally have a firm muscle tone and extremely sensitive stretch receptors. As a result, very fine adjustments are continually being made, and you are not aware of the cycles of contraction and relaxation that occur. Stretch reflexes are only one type of postural reflex; there are many complex polysynaptic postural reflexes. The CNS can adjust the sensitivity of muscle spindles by stimulating gamma motor neurons. This ability is important whenever voluntary contractions change the length of a muscle. For example, suppose you partially flex your elbow by contracting the biceps brachii muscle. You would expect that, in the new position, the muscle spindles would be compressed and muscle tone decreased.

That is not the case, because impulses arriving over gamma efferents cause the contraction of myofibrils in the intrafusal fibers in the biceps brachii muscle. The myofibrils pull on the membrane in the sensory portion of the intrafusal fiber until that membrane is stretched to its normal resting length. As a result, the muscle spindles remain sensitive to any externally imposed changes in muscle length. For instance, if someone drops a ball into your palm with your arm in this position, the muscle spindles will automatically adjust muscle tone to compensate for the increased load.

POLYSYNAPTIC REFLEXES

Polysynaptic reflexes can produce far more complicated responses than can monosynaptic reflexes. One reason is that the interneurons involved can control several muscle groups. Moreover, these interneurons may produce either excitatory or inhibitory postsynaptic potentials (EPSPs or IPSPs) at CNS motor nuclei, so the response can involve the stimulation of some muscles and the inhibition of others.

The Tendon Reflex

The stretch reflex regulates the length of a skeletal muscle. The tendon reflex monitors the external tension produced during a muscular contraction and prevents tearing or breaking of the tendons. The sensory receptors for this reflex have not been identified, but they are distinct from both muscle spindles and Golgi tendon organs, proprioceptors in tendons. The receptor is stimulated when the collagen fibers are stretched to a dangerous degree. In the spinal cord, these neurons stimulate inhibitory interneurons that innervate the motor neurons controlling the skeletal muscle. The greater the tension in the tendon, the greater the inhibitory effect on the motor neurons. As a result, skeletal muscles generally cannot develop enough tension to break their tendons.

Withdrawal Reflexes

Withdrawal reflexes move affected parts of the body away from a source of stimulation. The strongest withdrawal reflexes are triggered by painful stimuli, but these reflexes are sometimes initiated by the stimulation of touch receptors or pressure receptors. The flexor reflex, a representative withdrawal reflex, affects the muscles of a limb. When you step on a tack, a dramatic flexor reflex is produced in the affected leg. When the pain receptors in your foot are stimulated, the sensory neurons activate interneurons in the spinal cord that stimulate motor neurons in the anterior gray horns. The result is a contraction of flexor muscles that yanks your foot off the ground.

When a specific muscle contracts, opposing muscles must relax to permit the movement. For example, the flexor muscles that bend the leg are opposed by extensor muscles that straighten it out. A potential conflict exists: In theory, the contraction of a flexor muscle should trigger a stretch reflex in the extensors that would cause them to contract, opposing the movement. Interneurons in the spinal cord prevent such competition through reciprocal inhibition. When one set of motor neurons is stimulated, those neurons that control antagonistic muscles are inhibited. The term reciprocal refers to the fact that the system works both ways. When the flexors contract, the extensors relax; when the extensors contract, the flexors relax.

Withdrawal reflexes are much more complex than any monosynaptic reflex. They also show tremendous versatility, because the sensory neurons activate many pools of interneurons. If the stimuli are strong, interneurons will carry excitatory and inhibitory impulses up and down the spinal cord, affecting motor neurons in many segments. The end result is always the same: a coordinated movement away from the stimulus. But the distribution of the effects and the strength and character of the motor responses depend on the intensity and location of the stimulus.

When you step on something sharp, mild discomfort might provoke a brief contraction in muscles of your ankle and foot. More-powerful stimuli would produce coordinated muscular contractions affecting the positions of your ankle, foot, and leg. Severe pain would also stimulate contractions of your shoulder, trunk, and arm muscles. These contractions could persist for several seconds owing to the activation of reverberating circuits. In contrast, monosynaptic reflexes are relatively invariable and brief; the patellar reflex is completed in roughly 20 msec.

Crossed Extensor Reflexes

The stretch, tendon, and withdrawal reflexes are ipsilateral reflex arcs, in which the sensory stimulus and the motor response occur on the same side of the body. The crossed extensor reflex is called a contralateral reflex arc, because the motor response occurs on the side opposite the stimulus. The crossed extensor reflex complements the flexor reflex, and the two occur simultaneously.

When you step on a tack, while the flexor reflex pulls that foot away from the ground, the crossed extensor reflex stiffens your other leg to support your body weight. In the crossed extensor reflex, the axons of interneurons responding to the pain cross to the other side of the spinal cord and stimulate motor neurons that control the extensor muscles of the uninjured leg. Your opposite leg straightens to support the shifting weight. Reverberating circuits use positive feedback to ensure that the movement lasts long enough to be effective, despite the absence of motor commands from higher centers of the brain.

Polysynaptic reflexes range in complexity from a simple tendon reflex to the complex and variable reflexes associated with standing, walking, and running. Yet all polysynaptic reflexes share the same five basic characteristics: They involve pools of interneurons. Processing occurs in pools of interneurons before motor neurons are activated. The result may be excitation or inhibition; the tendon reflex produces inhibition of motor neurons, whereas the flexor and crossed extensor reflexes direct specific muscle contractions. They are intersegmental in distribution. The interneuron pools extend across spinal segments and may activate muscle groups in many parts of the body.

They involve reciprocal inhibition. Reciprocal inhibition coordinates muscular contractions and reduces resistance to movement. In the flexor and crossed extensor reflexes, the contraction of one muscle group is associated with the inhibition of opposing muscles. They have reverberating circuits, which prolong the reflexive motor response. Positive feedback between interneurons that innervate motor neurons and the processing pool maintains the stimulation even after the initial stimulus has faded. Several reflexes may cooperate to produce a coordinated, controlled response. As a reflex movement is under way, any antagonistic reflexes are inhibited. For example, during the stretch reflex, antagonistic muscles are inhibited; in the tendon reflex, antagonistic muscles are stimulated. In complex polysynaptic reflexes, commands may be distributed along the length of the spinal cord, producing a well-coordinated response.

INTEGRATION & CONTROL OF SPINAL REFLEXES

Reflex motor behaviors occur automatically, without instructions from higher centers. However, higher centers can have a profound effect on reflex performance. Processing centers in the brain can facilitate or inhibit reflex motor patterns based in the spinal cord. Descending tracts originating in the brain synapse on interneurons and motor neurons throughout the spinal cord. These synapses are continuously active, producing EPSPs or IPSPs at the postsynaptic membrane.

VOLUNTARY MOVEMENTS AND REFLEX MOTOR PATTERNS

Centers in the brain can interact with the stereotyped motor patterns programmed into the spinal cord. By making use of preexisting patterns, relatively few descending fibers can control complex motor functions. For example, the motor patterns for walking, running, and jumping are primarily directed by neuronal pools in the spinal cord. The descending pathways provide appropriate facilitation, inhibition, or "fine-tuning" of the established patterns. When complicated voluntary movements are under way, the neurons involved with spinal reflexes assist with muscular coordination and control.

For instance, the descending tracts that stimulate motor neurons controlling the biceps brachii muscle, brachialis muscle, and other elbow flexors also stimulate (1) the gamma efferents to the intrafusal fibers in these muscles and (2) inhibitory interneurons to relax antagonistic muscle groups, such as the triceps brachii muscle. The gamma efferents regulate the sensitivity of the stretch reflex, and the interneurons are the same as those activated in the withdrawal and crossed extensor reflexes. As muscle contraction proceeds, tendon reflexes keep tension production within tolerable limits. Motor control therefore involves a series of interacting levels. At the lowest level are monosynaptic reflexes that are rapid but stereotyped and relatively inflexible. At the highest level are centers in the brain that can modulate or build on reflexive motor patterns.