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SECTION F The Sensory System

CHAPTER
31
Overview of the Sensory System

he sensory system places the individual in Sensory systems may function on a conscious
T relationship to the environment. Every sen-
sation depends on impulses that arise by
or unconscious level. Unconscious visceral sensory
systems help regulate the internal environment. The
stimulation of receptors or end organs. These impulses monitoring of limb position in space has both a
are carried to the central nervous system (CNS) by conscious component—the posterior column path-
sensory nerves and then conveyed through fiber tracts ways—and an unconscious component—the spi-
to higher centers for conscious recognition, reflex nocerebellar pathways. The conscious somatosensory
action, or other consequences of sensory stimulation. system has two components: the position/vibration/
Somatic sensation is all senses other than the special fine discriminatory touch system and the pain/
senses. In this section, only general somatic sensory temperature/crude touch system. The different sen-
modalities are considered; the special senses—smell, sory modalities are carried over nerve fibers that vary
vision, taste, hearing, and vestibular sensation—are in size, diameter, and myelination. Sensory impulses
discussed with the cranial nerves that mediate them. are carried to the dorsal (posterior) root ganglia and
The sensory system can be classified in several dif- then into the CNS. After one or more synapses, the
ferent ways. Sherrington divided sensation into extero- impulses ascend specific fiber tracts and reach the
ceptive, interoceptive, and proprioceptive. Exteroceptive central sensory areas of the brain. Fine touch, posi-
sensation provides information about the external tion, and vibration from the body are carried over the
environment, including somatosensory functions and posterior column/medial lemniscus system. These
special senses. The interoceptive system conveys infor- sensations from the head and face are processed by the
mation about internal functions, blood pressure, or the trigeminal principal sensory nucleus in the pons. Pain
concentration of chemical constituents in bodily flu- and temperature from the body is carried over the
ids. Proprioception senses the orientation of the limbs spinothalamic tracts and from the head and face over
and body in space. Anatomists differentiate between the spinal tract and nucleus of the trigeminal. The
somatic and visceral sensation, with general and special major sensory pathways are depicted in Figure 31.1.
varieties of each. General somatic afferent fibers carry
exteroceptive and proprioceptive information; general
visceral afferent fibers carry impulses from visceral struc- SENSORY RECEPTORS
tures. Special somatic afferent fibers subserve the special
senses; special visceral afferent fibers mediate smell and The interface between the sensory nervous system
taste. Other terms used to categorize types of sensation, and the environment is the receptor. There are many
such as epicritic, protopathic, vital, and gnostic are of different types of receptors in the skin, subcutaneous
historical interest but have fallen into disuse. tissues, muscles, tendons, periosteum, and visceral
513

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514 SECTION F | THE SENSORY SYSTEM

Sensory association cortex

Primary
somesthetic
cortex

Thalamic
radiations Thalamus
VPL VPM

Trigeminothalamic Trigeminal principal


tract sensory nucleus

Medial lemniscus

Spinal tract of V
Nucleus of the spinal tract
Nucleus gracilis and cuneatus

Spinothalamic
tract Midline

Posterior columns

Posterior horn

Anterior commissure

FIGURE 31.1 The light touch, pressure, position, and vibration pathways from the body and face are indicated by the dashed line;
the pain and temperature fibers from the body and face are indicated by the solid line. Fibers from these various sources ultimately
converge on the ventral posterior nuclei of the thalamus, which projects via the thalamic radiations to the primary sensory cortex in
the postcentral gyrus. V, trigeminal; VPL, ventral posterior lateral; VPM, ventral posterior medial.

structures to subserve the transduction of various types varying degrees. Some receptors are rapidly adapting
of sensory information into nerve impulses. Sensory and most sensitive to on-and-off stimuli. Others adapt
end organs are found in the skin and mucous mem- slowly and function to constantly monitor a stimulus.
branes throughout the body. They are denser on the Receptors are the terminal part of, and are continu-
tongue, lips, genitalia, and fingertips and farther apart ous with, a sensory nerve. Receptor potentials induce
on the upper arms, buttocks, and trunk. One nerve action potentials in the nerve, with the frequency of
fiber may innervate more than one receptor, and each the action potential discharge usually in proportion
end organ may receive filaments from more than one to the amplitude of the receptor potential, which is
nerve fiber. Receptors may respond to more than one in turn proportional to the intensity of the applied
type of stimulus but have “specificity” because their stimulus. Each neuron has a specific receptive field,
threshold is lowest for a particular type of stimulus. which consists of all the receptors it can respond to.
Receptor stimulation causes a change in the perme- The receptive fields form more or less discrete maps
ability of its membrane that gives rise to a receptor or in the nervous system in which specific regions of the
generator potential—a local, nonpropagated poten- body are represented in specific regions of the brain.
tial whose intensity is proportional to the intensity Some systems have a highly organized map (e.g., the
of the stimulus. Receptors may adapt to a stimulus to somatosensory homunculus in the postcentral gyrus).

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CHAPTER 31 | OVERVIEW OF THE SENSORY SYSTEM 515

In other systems, the maps are crude. In the cortex, as mechanoreceptors. In encapsulated nerve endings,
neurons subserving the same modality and with simi- nonneural cells form a capsule around the terminal
lar receptive fields are organized into vertical rows, axon. Examples include Golgi tendon organs, muscle
which extend from the cortical surface to the white spindles, Ruffini endings, peritrichial endings, and
matter and are referred to as cortical columns. Meissner’s and Pacinian corpuscles.
Receptors may be free nerve endings (FNE), or There is evidence that abnormalities may be lim-
they may be encapsulated or connected to specialized ited to sensory receptors in some neuropathies previ-
nonneural components to form the sense organ. The ously thought to selectively affect small nerve fibers.
nonneural elements are not excitable, but they help to
form a structure that efficiently stimulates and excites
the sensory nerve fiber. Exteroceptors respond to exter- NERVE FIBER CLASSIFICATION
nal stimuli and lie at or near the interfaces between
the body and the environment. Special sensory extero- In the peripheral nervous system, axons are divided
ceptors subserve vision, hearing, smell, taste, and ves- into three major size groups: large myelinated, small
tibular function. General or cutaneous sensory organs myelinated, and unmyelinated. The largest fibers are
include the free and encapsulated receptor terminals spindle afferents and motor fibers arising from alpha
in the skin. Proprioceptors respond to stimulation motor neurons. The smallest, unmyelinated fibers
of deeper tissues, such as muscles and tendons, and are pain and postganglionic autonomic fibers. Large
are designed particularly to detect movement and the myelinated axons have diameters in the 6 to 12 mm
position of body parts. Receptors around hair follicles range, small myelinated axons 2 to 6 mm, and unmy-
are activated by distortion of the hairs. elinated axons 0.2 to 2 mm. Small myelinated fib-
Receptors may be classified by the specific ers are about three times more numerous than large
modality to which they are more responsive, such as myelinated axons. The conduction velocity (CV) of a
mechanoreceptors, thermoreceptors, chemoreceptors, fiber depends on its diameter and degree of myelina-
photoreceptors, and osmoreceptors. Mechanoreceptors tion. Large fibers conduct more rapidly than small
respond to deformation, such as touch or pressure. ones, and myelinated fibers, more rapidly than unmy-
Stimulation of mechanoreceptors causes a physical elinated ones. The CV ranges from less than 1 m/s for
deformation of the receptor that results in the open- small, unmyelinated fibers to greater than 100 m/s
ing of ion channels. Polymodal receptors respond effi- for large, myelinated fibers. In large, myelinated fib-
ciently to more than one modality, especially stimuli ers, the fiber diameter (in mm) × 6 approximates the
that cause tissue damage and pain. There is a great CV (in m/s).
deal of variation in the density of sensory receptors Peripheral nerve fibers are classified by size
between different body surface regions. Also, receptor and CV according to two schemes: the ABC and
density decreases with advancing age. the I/II/III/IV systems (see Chapter 23). The ABC
Receptors may also be classified morphologically, scheme includes both motor and sensory fibers. The
but the correlation between function and morphol- A-alpha and A-gamma fibers are motor. The A-alpha
ogy is not nearly as close as was once believed. There group also includes afferents from encapsulated recep-
are FNE, epidermal endings, and encapsulated end- tors in the skin, joints, and muscles, including the pri-
ings. The FNE are fine, unmyelinated terminal fibers mary spindle afferents. The A-beta and A-delta fibers
that arborize in the skin, fascia, ligaments, tendons, are primarily cutaneous afferents. Group B fibers are
and other connective tissues throughout the body. preganglionic autonomics. Group C fibers include
They mediate several sensory modalities; some are postganglionic autonomics, general visceral afferents,
exclusively nociceptors. The FNE are the terminals and pain and temperature fibers. The I/II/III/IV sys-
of sensory C fibers or A-delta fibers (see “Nerve Fiber tem applies only to afferent fibers. Groups I to III
Classification”) and are located in both glabrous and are myelinated; group IV is unmyelinated. The Ia fib-
hairy skin. The FNE terminals of unmyelinated nerve ers are spindle afferents from nuclear bag fibers; the
fibers are mainly nociceptive, but they may also be Ib fibers arise from Golgi tendon organs; and the II
thermoreceptors or mechanoreceptors. Merkel cell fibers are spindle afferents from nuclear chain fibers.
endings (tactile discs or menisci) are specialized nerve Group III fibers are cutaneous axons approximately
endings lying just below the epidermis, especially in the same as A-delta fibers. Group IV fibers corre-
glabrous skin, and around hair follicles that function spond to C fibers and are primarily nociceptive.

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516 SECTION F | THE SENSORY SYSTEM

In addition to the relationships between nerve those to light touch. Figure 36.5 shows the dermat-
fiber diameter, CV and sensory modality, the vulner- ome distributions as depicted by Keegan and Garrett.
ability to various types of injury varies with size and
type of fiber. Cocaine, which blocks the conduction
of the smaller fibers first, causes loss of sensation in ANATOMY OF THE POSTERIOR ROOT
the order of slow pain, cold, warmth, fast pain, touch,
and position. Pressure, which blocks the conduction The oval-shaped dorsal root ganglia (DRG) lie on the
of the larger fibers first, causes loss of sensation in posterior root in the intervertebral foramen, just lat-
the order of position, vibration, pressure, touch, fast eral to the point where the posterior root penetrates
pain, cold, warmth, and slow pain. Most peripheral the dura. The connective tissue capsule around each
neuropathies affect both large and small fibers, but DRG is continuous with the epineurium of the spi-
in some conditions, the involvement primarily affects nal root. The DRG is composed of neurons, satel-
either the large or the small fibers. lite cells, and a highly vascular supporting stroma.
The DRG neurons are unipolar. A single nonmy-
elinated “dendro-axonal” process leaves the cell and
then bifurcates into peripheral and central branches.
DERMATOMES
The peripheral processes conduct afferent impulses
Sensory nerve roots supply cutaneous innervation to toward the cell body; they are functionally elongated
specific dermatomes. The dermatome innervation of dendrites but more closely resemble axons from a
the extremities is complex, in part due to the migration structural standpoint and by convention are referred
of the limb buds during embryonic development. As to as axons. Large sensory neurons may be found sin-
a result, the C4-C5 dermatomes abut T1-T2 on the gly or in small groups proximal or distal to the DRG.
upper chest, and the L1-L2 dermatomes are close to Sometimes, the entire DRG lies in an ectopic
the sacral dermatomes on the inner aspect of the thigh intraspinal location, well proximal to its usual posi-
near the genitalia. The generally available dermatomal tion, making it vulnerable to involvement by her-
charts are primarily derived from three sources: Head niated nucleus pulposus or osteophytic spur. Such
and Campbell, Foerster, and Keegan and Garrett, ectopic DRGs have been mistaken for tumors, with
who all used very different approaches. Head and unfortunate results. The DRG for the C1 posterior
Campbell were primarily interested in herpes zoster root is often missing.
and mapped dermatomes according to the distribu- The dorsal root is divided into a medial zone,
tion of herpetic eruptions. Foerster performed pos- conveying large fiber proprioceptive traffic and a lat-
terior rhizotomies in patients with chronic pain. He eral zone conveying small fiber pain and temperature
mapped the distribution of an intact root when one traffic. As the posterior root exits the DRG to enter
or more of those above and below had been severed the spinal cord, two discrete fascicles may be visible;
or by electrically stimulating the stump of a severed these correspond to the medial and lateral divisions.
root and observing the area of cutaneous vasodilation. After the posterior root joins the spinal cord, the
The observation of dermatomal overlap originated pathways serving different sensory modalities diverge
partly from this work, and for a time, many believed and follow very different central courses through the
a lesion of a single root would produce no detectable spinal cord and lower brainstem, only to draw closer
deficit. Keegan and Garrett examined a large series together as they ascend through the upper brainstem
of patients with clinical involvement of various roots to ultimately reconverge as they enter the thalamus.
and mapped the sensory deficits; there was surgical
correlation in 53% of the patients. The loss of sen-
sation due to isolated involvement of a single root, CLINICAL EXAMINATION
as occurs clinically, produces a different dermatomal
map than the preserved sensation in a zone of anes- Sensory function is divided clinically into primary
thesia as found by Foerster. It is clear that the derma- modalities and secondary or cortical modalities. The
tomal overlap is such that the clinical deficit from an primary modalities include touch, pressure, pain,
isolated root lesion is typically much more restricted temperature, joint position sense, and vibration. The
than that expected from the anatomical geography of cortical or secondary modalities are those that require
the dermatome. Deficits to pin prick are smaller than synthesis and interpretation of primary modalities

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CHAPTER 31 | OVERVIEW OF THE SENSORY SYSTEM 517

TABLE 31.1 Generally Accepted Definitions of Commonly Used Terms Regarding the Sensory System and
Abnormalities of Sensation
Term Definition

Allodynia Increase in sensibility to pain; pain in response to a stimulus not normally painful
Alloesthesia (allesthesia) Perception of a sensory stimulus at a site other than where it was delivered; tactile allesthesia is feeling
something other than at the site of the stimulus; visual allesthesia is seeing something other than where
it actually is (see mitempfindung, Box 31.1)
Analgesia (alganesthesia) Absence of sensibility to pain
Astereognosis Absence of spatial tactile sensibility; inability to identify objects by feel
Anesthesia Absence of all sensation
Dysesthesias Unpleasant or painful abnormal perverted sensations, either spontaneous or after a normally nonpainful
stimulus (e.g., burning in response to touch); often accompany paresthesias
Hypalgesia Decrease in sensibility to pain
Hyperalgesia Increase in sensibility to pain; pain in response to a stimulus not normally painful
Hyperpathia Increase in sensibility to pain; pain in response to a stimulus not normally painful
Kinesthesia The sense of movement
Pallesthesia Vibratory sensation (decreased, hypopallesthesia; absent, apallesthesia)
Paresthesias Abnormal spontaneous sensations experienced in the absence of specific stimulation (feelings of cold,
warmth, numbness, tingling, burning, prickling, crawling, heaviness, compression, or itching)

by the sensory association area in the parietal lobe. definition of esthesia is perception, feeling, or sen-
These include two-point discrimination, stereog- sation (Gr. aesthesis “sensation”). Algesia refers to
nosis, graphesthesia, tactile localization, and others. the sense of pain (Gr. algos “pain”). Hypalgesia is a
When the primary modalities are normal in a par- decrease, and analgesia (or analgesthesia) an absence,
ticular body region, but the cortical modalities are of pain sensation. The combining form “algia” refers
impaired, a parietal lobe lesion may be responsible. to any painful condition. Hypesthesia is a decrease,
Itch and tickle sensations are closely allied to pain; and anesthesia an absence, of all sensation. Paresthesia
they are probably perceived by the same nerve end- is an abnormal sensation; dysesthesia (Gr. dys “bad”)
ings and are absent following procedures used for the is an abnormal, unpleasant, or painful sensation.
relief of pain. Table 31.1 summarizes some of the definitions.
Many terms have been used, not always Seldom-used terms and those of primarily historical
consistently, to describe sensory abnormalities. The interest are summarized briefly in Box 31.1.

BOX 31.1
Other Sensory Terms
Anaphia, absence of sensibility to touch; arthresthe- myesthesia, muscle sensation, sensibility coming
sia, the perception of joint movement and position; from muscles; pallesthesia, vibratory sensation;
baresthesia, ability to sense pressure or weight; mitempfindung, distant referral of cutaneous sen-
barognosis, the appreciation, recognition, and sation; pallanesthesia, loss of vibratory perception;
differentiation of weight, the ability to differentiate piesesthesia, pressure sensibility; thermanesthesia,
between weights; abaragnosis, loss of ability to dif- loss of thermal sensibility; thermhyperesthesia,
ferentiate weight; bathyesthesia, deep sensibility, increase in thermal sensibility; thermhypesthe-
from parts of the body below the surface, such as sia, decrease in thermal sensibility; thigmesthesia,
muscles and joints; gargalanesthesia, absence of the light touch or general tactile sensibility; topesthesia
sensation of tickling; statognosis, the awareness of (topognosia), fine discriminatory and localized tac-
posture; isothermagnosia, perception of either cold tile sensibility, the ability to localize a tactile sensa-
or warm stimuli as warm—may be seen following tion; topoanesthesia (topagnosia), the inability to
cordotomy or with high spinal cord lesions; localize a tactile sensation.

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518 SECTION F | THE SENSORY SYSTEM

Sensory abnormalities may be characterized by understands the expected responses. During the
an increase, decrease, absence, or perversion of sensa- examination, the patient should be warm, comfort-
tion. An example of increased sensation is pain—an able, and relaxed. The best results are obtained when
unpleasant or disagreeable feeling that results from the patient is lying comfortably in a warm, quiet
excessive stimulation of certain sense organs, fibers, room. Obtaining patient confidence is important.
or tracts. Perversions of sensation take the form Satisfactory results cannot be obtained when the
of paresthesias, dysesthesias, and phantom sensa- patient is suspicious, in pain, uncomfortable, fearful,
tions. Impairment and loss of sensation result from confused, or distracted by sensations such as noise or
decreased acuity of the sensory organs or receptors, hunger. If the patient is in pain or discomfort, or if he
impaired conduction in sensory fibers or tracts, or has recently been sedated, the examination should be
dysfunction of higher centers causing impairment in postponed. The areas under examination should be
the powers of perception or recognition. uncovered, but it is best to expose the various parts of
The sensory examination is performed to dis- the body as little as possible. The patient’s eyes should
cover whether areas of absent, decreased, exaggerated, be closed or the areas under examination shielded to
or perverted sensation are present, and to determine eliminate distractions and to avoid misinterpretation
the type of sensation affected, the degree of abnormal- of stimuli. Homologous areas of the body should be
ity, and the distribution of the abnormality. Findings compared whenever possible.
may include loss, decrease, or increase of one or more The detail and technique used for the sensory
types of sensation; dissociation of sensation with loss examination depend on the history. For example, a
of one modality type but not of others; loss of ability patient with no sensory complaints referred for evalu-
to recognize differences in degrees of sensation; mis- ation of headache or vertigo requires only a screening
interpretations (perversions) of sensation; or areas of examination. A patient who is seen for possible carpal
localized hyperesthesia. More than one of these may tunnel syndrome, radiculopathy, peripheral neuropa-
occur simultaneously. thy, or a suspected parietal lobe lesion requires a very
The sensory examination is arguably the most different approach.
difficult and tedious part of the neurologic examina- The examiner should first determine whether
tion. Some examiners prefer to assess sensory func- the patient is aware of subjective changes in sensation
tions early in the course of the examination, when or is experiencing abnormal spontaneous sensations.
the patient is most likely to be alert and attentive. Sensory symptoms may be divided into negative
Fatigue causes faulty attention and slowing of the symptoms, lack of sensation, and positive symptoms,
reaction time, and the findings are less reliable when abnormal sensory discharges such as paresthesias
the patient has become weary during the examina- and dysesthesias. Positive and negative symptoms
tion. Others argue the routine sensory examination is may occur together. Inquire whether the patient has
the most subjective and least useful part of the neu- noticed pain, paresthesias, or loss of feeling; whether
rologic examination and prefer to leave it until the any part of the body feels numb, dead, hot, or cold;
end. Since the results depend largely on subjective whether he has perceived sensations such as tingling,
responses, the full cooperation of the patient is neces- burning, itching, “pins and needles,” pressure, disten-
sary if conclusions are to be accurate. Occasionally, tion, formication, or feelings of weight or constric-
objective evidence, such as withdrawal of the part tion. If such symptoms are present, determine their
stimulated, wincing, blinking, and changes in coun- type and character, intensity, distribution, duration,
tenance, may aid in the delineation of areas of sensory and periodicity, as well as exacerbating and relieving
change. Pupillary dilation, tachycardia, and perspira- factors. Spontaneous pain must be differentiated from
tion may accompany painful stimulation. Keenness tenderness. Pain and numbness may exist together,
of perception and interpretation of stimuli differ in as in thalamic pain and peripheral neuropathy. The
individuals, in various parts of the body, and in the patient’s manner of describing the pain or sensory
same individual under different circumstances. disturbance and the associated affective responses,
For a reliable sensory examination, the patient the nature of the terms used, the localization, and the
must understand the procedure and be ready and precipitating and relieving factors may aid in differen-
willing to cooperate. Accurate communication is tiating between organic and nonorganic disturbances.
vital. The purpose and method of testing should Nonorganic abnormalities are often associated with
be explained in simple terms, so that the patient inappropriate affect (either excessive emotionality or

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CHAPTER 31 | OVERVIEW OF THE SENSORY SYSTEM 519

indifference), are often vague in character or location, Accuracy in localization of pain, temperature,
and reactions to them are not consistent with the and tactile stimuli is also informative. Tactile locali-
degree of disability. zation is a sensitive test of sensory function; there
If the patient has no sensory symptoms, testing may be loss of localization before there is a detect-
can be done rapidly, bearing in mind the major sen- able change in sensory threshold. Tactile localization
sory nerve and segmental supply to the face, trunk, is most accurate on the palmar surfaces of the fingers,
and extremities. In certain situations, more careful especially the thumb and index finger. The patient
sensory testing is required. If there are specific sen- should name or point to the area stimulated, compar-
sory symptoms—motor symptoms such as atrophy, ing responses on the two sides of the body.
weakness, or ataxia—if any areas of sensory abnor- The results of the sensory examination may
mality are detected on the survey examination, or if at times seem unreliable and confusing. The pro-
the clinical situation suggests the likelihood of sen- cess can become tedious, and the findings difficult
sory abnormalities, then detailed sensory examina- to interpret. Sensory changes due to suggestion are
tion should be performed. The presence of trophic notoriously frequent in emotionally labile individu-
changes, especially painless ulcers and blisters, is also als, but suggestion can produce nonorganic findings
an indication for careful sensory testing since these in patients with organic disease. Care must be taken
may be the first manifestations of a sensory disorder in drawing conclusions. To obtain reliable results, it
of which the patient is unaware. In patients with lim- may be necessary to postpone the sensory examina-
ited cooperation, it may be desirable to examine the tion if the patient has become fatigued, or to repeat
areas of sensory complaint first and then survey the the testing at a later time. The sensory examination
rest of the body. should always be repeated at least once to confirm the
The simpler the method of examination, the findings. Sensory testing, more than any other part
more satisfactory the conclusions. Explain to the of the neurologic examination, requires patience and
patient what is to be done and demonstrate in an area detailed observation for reliable interpretation.
expected to be normal what the stimulus feels like. The following are some of the difficulties that
Then, have the patient close his eyes and begin the may be encountered in performing the sensory
testing. The subject should be asked to tell the type examination. The uncooperative patient may be
of stimulus perceived and its location, with the exam- indifferent to the sensory examination or object to
iner taking care not to suggest responses. Responses the use of painful stimuli. The overly cooperative
are normally prompt, and a consistent delay in patient, on the other hand, may make too much
answering may indicate an abnormal delay in percep- of small differences and report changes that are not
tion. There are two general screening patterns: side to present. Some areas of the body, such as the ante-
side and distal to proximal. The side-to-side screen- cubital fossae, the supraclavicular fossae, and the
ing should usually compare the major dermatomes neck, are more sensitive than others; apparent sen-
and peripheral nerve distributions, although more sory changes in these regions may lead to fallacious
abbreviated screening may be appropriate in certain conclusions. The last in a series of identical stimuli
clinical circumstances. Distal to proximal testing is may be interpreted as the strongest. Even though
appropriate when peripheral neuropathy is part of pain sensibility is absent, a patient may still be able
the differential diagnosis. The distribution of abnor- to identify a sharp stimulus with a pin. Occasionally
malities can be drawn on the skin with a marker and in syringomyelia, with lost pain but preserved tactile
recorded on a chart (Figure 36.5), indicating areas of sensibility, the patient may recognize the pin point
change in the various modalities by horizontal, ver- in an analgesic area and give confusing and incon-
tical, or diagonal lines, stippling or different colors. sistent responses. Sensory findings are difficult to
A key helps to explain the meaning of the various evaluate in individuals with low intellectual endow-
symbols and colors, as does a note regarding the coop- ment, language difficulties, or a clouded sensorium,
eration and insight of the patient and an estimate of but it may be necessary to carry out the examina-
the reliability of the examination. Sensory charts are tion despite these obstacles. In patients with altered
helpful for comparison with the results of subsequent mental status or a decreased sensorium, pain may
examinations in following the course of the patient’s be tested grossly by pricking or pinching the skin,
illness, and for comparison with the results of other comparing responses on the two sides of the body. In
examiners. such patients, it may only be possible to determine

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520 SECTION F | THE SENSORY SYSTEM

whether or not the patient reacts to painful stimuli Dyck PJ, Zimmerman I, Gillen DA, et al. Cool, warm, and
heat-pain detection thresholds: testing methods and infer-
in various parts of the body. A child may be fearful
ences about anatomic distribution of receptors. Neurology
of testing, requiring assurance at the outset that the 1993;43:1500–1508.
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In young children, it is often best to delay sensory rology. Semin Neurol 2002;22:399–408.
Gilman S, Newman SW. Manter and Gatz’s Essentials of Clinical
testing until the end of the examination, particularly Neuroanatomy and Neurophysiology. 10th ed. Philadelphia: FA
when even mildly uncomfortable, yet threatening, Davis, 2003.
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CHAPTER
32
The Exteroceptive Sensations

xteroceptive sensations originate in peripheral of neurotransmitters; pain impulses are thought to


E receptors in response to external stimuli and
changes in the environment. There are four
be mediated primarily by substance P and glutamate.
Activity in the spinothalamic tract (ST) neurons of
main types of general somatic sensation: pain, thermal the posterior horn is modulated by descending path-
or temperature sense, light touch or touch-pressure, ways. Stimulation of certain brain regions inhibits the
and position sense or proprioception. response of ST cells to noxious stimuli. Descending
influences are known to arise from the nucleus raphe
magnus, periaqueductal gray, brainstem reticular for-
mation, periventricular gray, ventral posterior lateral
PAIN AND TEMPERATURE (VPL) thalamic nucleus, and the parietal cortex and
SENSATION travel primarily in the corticospinal tract and dorso-
lateral funiculus. These pathways are important in
Anatomy and Physiology pain control mechanisms.
Impulses carrying superficial pain sensation arise in The majority of axons originating from second-
nociceptors—free or branched nerve endings in the order spinothalamic neurons cross the midline in
skin and mucous membranes. Some nociceptors the anterior white commissure and gather into the
respond to specific types of stimuli, whereas others anterior and lateral STs; a small proportion of fibers
are polymodal. Thermoreceptors for heat and cold ascend ipsilaterally. Fibers crossing in the anterior
sensation are free nerve endings in the dermis. Warm white commissure are affected early in syringomyelia.
and cold stimuli activate different fibers. Pain and In the past, anatomists thought the anterior ST car-
thermal sensation are carried along small myelinated ried crude touch and the lateral ST pain and temper-
A-delta and unmyelinated C nerve fibers to the dor- ature; current evidence suggests all these modalities
sal root ganglion (DRG), where the first cell body is are carried in both tracts, so the lateral and anterior
situated (Figure 32.1). The impulses in response to STs are now sometimes lumped together as the ante-
moderate heat or cold travel primarily over A-delta rolateral or ventrolateral system (ALS) or simply the
and some C fibers. The response to the pain associ- spinothalamic tract or system. For clinical purposes,
ated with the extremes of temperature is conveyed it remains useful to consider the pain and temper-
along C fibers. Axons from small and intermediate ature pathways in the ST as a distinct system. The
size neurons in the DRG traverse the lateral divi- ST ascends in an anterolateral position, just medial
sion of the dorsal root to enter the dorsolateral fas- to the anterior spinocerebellar tract (Figure 32.2).
ciculus of the spinal cord (Lissauer’s tract), where they Intermingled with the fibers of the ST are ascending
ramify longitudinally for one or two segments. The spinoreticulothalamic fibers, which contribute to the
axons leave Lissauer’s tract, enter the posterior gray ALS. The ST is somatotopically organized, and the
horn, and synapse in laminae I to V. Second-order distribution of fibers is clinically relevant. Lowermost,
neurons for the spinothalamic system lie primarily sacral and lumbar, fibers entering first are displaced
in laminae I, II, and V (see Chapter 24). The other progressively more laterally by subsequently entering
related posterior horn cells are interneurons in the fibers. As the tract ascends, the sacral fibers come to
pain pathway. The posterior horn contains a variety lie most lateral and superficial, nearer to the surface

521

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522 SECTION F | THE SENSORY SYSTEM

Pacinian
corpuscle
Medial division of dorsal root

Fasciculus gracilis

Fasciculus cuneatus
Muscle
spindle

Lateral
Lateral spinothalamic
Meissner's division of tract
tactile dorsal root
Anterior
corpuscle Anterior horn Anterolateral
spinothalamic
motor neuron system
tract

Free sensory
ending
FIGURE 32.1 Diagram of the spinal cord and dorsal root showing the peripheral receptors and terminations of fibers within the
spinal cord.

of the cord (Figure 32.3), with cervical fibers most lateral to the medial lemniscus (ML) and medial to
medial. There is also a slight rotation so that the sacral the middle cerebellar peduncle; in the mesencepha-
fibers also come to lie somewhat more posterior as lon, it is peripheral, dorsal to the ML and just dorso-
the tract ascends. At midbrain levels, lower-extremity lateral to the red nucleus. It passes near the colliculi
and sacral fibers are posterior, and those from the and enters the diencephalon just medial to the bra-
upper limb and trunk are more anterior. Since the chium of the inferior colliculus.
sacral fibers lie most laterally, an intramedullary spi- Pain and temperature fibers from the face enter
nal cord lesion, such as a neoplasm, may produce the pons through the Gasserian ganglion and then
“sacral sparing,” preservation of sensation in a saddle descend in the spinal tract of the trigeminal nerve to
distribution in the face of sensory loss otherwise pre- varying levels, where they synapse on neurons in the
sent below a certain spinal level. Conversely, a com- adjacent nucleus of the spinal tract (see Chapter 15).
pressive lesion pressing on the upper spinal cord may These second-order neurons decussate and form the
preferentially involve the sacral spinothalamic fibers, trigeminothalamic tract, which runs near the ascend-
causing sacral dysfunction first. Fibers carrying deep ing spinothalamic and lemniscal fibers (Figure 15.2).
pain are in general thought to lie nearer the midline The other cranial nerves carrying exteroceptive pain
than those carrying superficial pain. The spinoreticu- sensation have ganglia comparable to the DRG and
lothalamic fibers in the ALS subserve diffuse, poorly pathways corresponding to the trigeminothalamic
localized pain from deep and visceral structures. They tract. These are discussed in the chapters on the indi-
may also be involved in the affective aspects of pain. vidual cranial nerves.
In the medulla, the ST lies peripherally, dorso- In the upper lateral midbrain, all the somatosen-
lateral to the inferior olivary nuclei; in the pons, it is sory fibers begin to converge. The ST fibers are joined

Campbell_Chap32.indd 522 6/21/2012 5:03:40 PM


Nuclei ventralis
posterolateralis (VPL) and
posteromedialis (VPM) of thalamus

Gasserian (trigeminal)
ganglion

FIGURE 32.2 The lateral spinothalamic tract.

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524 SECTION F | THE SENSORY SYSTEM

FIGURE 32.3 Diagram of cross section of the cervical region of the spinal cord showing the arrangement of fibers in the spinotha-
lamic and pyramidal tracts and dorsal columns. Heavy dots indicate fibers carrying temperature sensation, crosses indicate fibers
carrying pain sensation, and fine dots indicate fibers carrying tactile impulses. C, T, L, and S indicate fibers from or destined for
cervical, thoracic, lumbar, and sacral levels of the spinal cord.

in the rostral brainstem by the laterally migrating the parietal lobe; those from the face terminate on
fibers from the ML and by ascending trigeminotha- the lateral, inferior portion of the postcentral gyrus
lamic fibers so that ultimately all the fibers subserving (Figure 6.7). Fibers of the spinoreticulothalamic tract
somatosensory function run together as they approach carry nociceptive information in the ALS. There are
the thalamus. The tracts enter the ventrobasal and synapses in the brainstem reticular formation and
ventral posterior nucleus of the thalamus together; medial part of the thalamus. Spinoreticulothalamic
body sensation fibers terminate in the VPL nucleus fibers terminate in the intralaminar thalamic nuclei.
and facial sensation fibers in the ventral posterior The thalamic neurons that mediate pain project
medial (VPM) nucleus. There is detailed somatotopic both to the parietal lobe and to the limbic cortex.
organization within VPL and VPM. From the thala- Projections from the intralaminar nuclei terminate
mus, fibers run in the thalamic radiations through the in the hypothalamus and limbic system and prob-
posterior limb of the internal capsule to the primary ably mediate the affective and autonomic responses
somesthetic cortex in the postcentral gyrus for con- to pain.
scious recognition. The primary somesthetic cortex Descending pathways serve to modulate pain.
communicates with the parietal sensory association Fibers from the frontal cortex and hypothalamus
cortex and with other cortical areas. Thalamocortical project to the midbrain periaqueductal gray. The
fibers also project to the superior bank of the sylvian descending pain modulation pathway then descends
fissure. in the dorsal part of the lateral funiculus to the poste-
In the thalamoparietal radiations, fibers carry- rior horn. Descending fibers from the locus ceruleus,
ing lower-extremity sensation curve medially to the the raphe nuclei and other brainstem areas also
superior medial surface of the hemisphere adjacent modulate the pain response. These descending path-
to the medial longitudinal fissure; those from the ways are important in endogenous pain control and
upper body go to the midportion of the surface of opiate analgesia.

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CHAPTER 32 | THE EXTEROCEPTIVE SENSATIONS 525

Clinical Examination for example, “If this (stimulating the apparently


normal side) side is a dollar’s worth (or 100%), how
There are many methods for testing superficial pain much is this (stimulating the apparently abnormal
sensation. A simple and commonly used method, as side) worth?” The overanalytical but neurologically
reliable as any, is to use a common safety pin bent normal patient often responds with an estimate
at right angles so its clasp may serve as a handle. on the order of “95 cents,” while the patient with
The instrument should be sharp enough to create a real, clinically significant sensory loss is more apt
mildly painful sensation, but not so sharp as to draw to respond with “5 cents” or “25 cents.” Delivering
blood. A hypodermic needle is far too sharp unless alternately sharp and dull stimuli, as with the sharp
its point has been well blunted against some hard and blunt ends of a safety pin and instructing the
surface. A broken wooden applicator stick is often patient to reply “sharp” or “dull” is frequently useful
used and is usually satisfactory provided the shards but may not detect subtle sensory loss only detect-
are sharp. Adequately sharp ends can be obtained by able in comparison with an uninvolved area. Slight
holding the stick at the very ends while breaking it. changes can sometimes be demonstrated in a coop-
Disposable sterile devices, sharp on one end and dull erative patient by asking her to indicate the altera-
on the other, are commercially available. While it is tions in sensation when a pinpoint is drawn lightly
not necessary for the stimulating instrument to be over the skin. A cooperative patient with a discrete
sterile, whatever is used must be discarded after use distribution of sensory loss may be able to map out
on a single patient to avoid the risk of transmitting the involved area quite nicely if instructed how to
disease from accidental skin puncture. There is no proceed and left alone for a short time with tools
place in modern neurology for reusable sharp instru- and a marking instrument. The affected area can
ments such as the Wartenberg wheel, but dispos- then be compared with a figure showing sensory
able pinwheels are available. Various sensory testing distributions.
devices have been used experimentally. Instruments The latent time in the response to stimulation
for evaluating sensation quantitatively are available is eliminated and the delineation more accurate if
commercially. the examination proceeds from areas of lesser sensi-
A helpful trick is to hold the pin or shaft of the tivity to those of greater sensitivity rather than the
applicator stick lightly between thumb and fingertip, reverse. If there is hypalgesia, move from areas of
and let the shaft slide between fingertip and thumb decreased sensation to those of normal sensation; if
tip with each stimulation. This helps insure more there is hyperalgesia, proceed from the normal to
consistent stimulus intensity than putting a fingertip the hyperalgesic area. There may be a definite line
on the end of the instrument and trying to control of demarcation between the areas of normal and
the force with the hand or wrist. Experience teaches abnormal sensation, a gradual change, or at times a
how to gauge the intensity of the applied stimulus zone of hyperesthesia between them. It is occasion-
and the expected reaction to it. The clinical evalu- ally useful to move from the normal to the numb
ation of superficial pain, temperature, and touch area. In myelopathy, a spinal sensory level that is the
sensation shows a reasonably good correlation with same going from rostral to caudal as from caudal to
quantitative assessment. rostral suggests a very focal and destructive lesion;
It is best to do the examination with the when the two levels are far apart, the lesion is usu-
patient’s eyes closed. The patient should be asked ally less severe. If testing is done too rapidly, the area
to judge whether the stimulus feels as sharp on one of sensory change may be misjudged. Applying the
side as on the other. Always suggest the stimuli stimuli too close together may produce spatial sum-
should be the same, as by language such as, “Does mation; stimulating too rapidly may produce tempo-
this feel about the same as that?” Avoid such lan- ral summation. Either of these may lead to spurious
guage as “Does this feel any different?” or “Which findings. If stimulation is too rapid, or if conduction
feels sharper?” Suggesting there should be a differ- is delayed, a given response may refer to a previous
ence encourages some patients to overanalyze and stimulation. Stimuli should be applied at irregular
predisposes them to spurious findings and a tedi- intervals to avoid patient anticipation. If the patient
ous, often unreliable examination. A commonly knows when to expect a stimulus, a seemingly nor-
used technique is asking the patient to compare one mal response can occur even from an anesthetic area.
side to the other in monetary or percentage terms, Include control stimuli from time to time, especially

Campbell_Chap32.indd 525 6/21/2012 5:03:47 PM


526 SECTION F | THE SENSORY SYSTEM

if the patient is comparing sharp and dull (e.g., using in patients with circulatory insufficiency or vasocon-
the dull end of the pin while asking if it is sharp), to striction causing acral coolness.
be sure the patient has understood the instructions Quantitative sensory testing (QST) uses neuro-
and is paying attention. physiologic methods to examine sensation. It provides
Temperature sensation may be tested with test very accurately measured stimuli of various types
tubes containing warm and cool water, or by using and uses strict paradigms for recording responses.
various objects with different thermal conductivity. Temperature sensation is tested by delivering pulses
Ideally, for testing cold, the stimuli should be 5°C of hot and cold and determining the threshold for
to 10°C (41°F to 50°F), and for warmth, 40°C to detection. Extremes of temperature assess pain. There
45°C (104°F to 113°F). The extremes of free-flow- is good correlation between QST and clinical meth-
ing tap water are usually about 10°C and 40°C. ods, but QST is very useful for longitudinal studies.
Temperatures much lower or higher than these elicit
pain rather than temperature sensations. Normally,
it is possible to detect a difference of about 1°C in
the range of around 30°C. The tubes must be dry, TACTILE SENSATION
as dampness may be interpreted as cold. The tines
of a tuning fork are naturally cool and work well for
Anatomy and Physiology
giving a quick impression of the ability to appreci- Cutaneous receptors that mediate light touch or
ate coolness. The tines quickly warm with repeated general tactile sensibility include free nerve end-
skin contact; applying the tines alternately and wav- ings, Merkel cell endings, and encapsulated end-
ing the fork in the air between stimuli helps prevent ings such as Meissner’s and Pacinian corpuscles
this warming. Holding the tines under cold running and Ruffini endings. All the encapsulated recep-
tap water may also be helpful. Some examiners warm tors function as mechanoreceptors with afferent
one tine deliberately by rubbing and then test the nerve fibers in the group II and III range. Pacinian
ability to discriminate between the warm side and corpuscles are large, lamellated structures located
the cool side of the fork. This technique has limited subcutaneously in the palmar, plantar and digital
practicality because the cool side warms so rapidly skin, genitalia, and other sensitive areas; they func-
with skin contact. The latency for detecting tem- tion as rapidly adapting mechanoreceptors. They
perature is longer than for other sensory modalities, are especially responsive to vibration, most notably
and the application of the stimulus may need to be in the 40 to 1,000 Hz frequency range. Meissner
extended. tactile corpuscles are found primarily in thick hair-
In the general examination, it is sufficient to less skin, such as the hand, foot and lips, and are
determine whether the patient can distinguish hot most highly developed in the finger pads. They also
and cold stimuli. It may be useful in some circum- respond to vibration in the low-frequency range
stances, such as the detection of mild peripheral neu- (10 to 400 Hz) and are maximally sensitive at 100
ropathy, to determine whether the patient is able to to 200 Hz. Merkel cell receptors are also slowly
differentiate between slight variations in temperature. adapting mechanoreceptors that respond to low fre-
This is best done with special devices for testing tem- quency vibration. Ruffini endings are slowly adapt-
perature sensation quantitatively. In most instances, ing mechanoreceptors located in hairy as well as
heat and cold sensibility are equally impaired. Rarely, glabrous skin, in joint capsules, tendon insertions,
one modality may be involved more than the other; and elsewhere. They are particularly responsive to
the area of impaired heat sensibility is usually the stretching or indentation of the skin.
larger. Pain and temperature sensibility are usually Light touch sensation is conveyed over large and
involved equally with lesions of the sensory system, small myelinated peripheral nerve fibers to unipolar
and it is rarely necessary to test both. Testing tem- DRG cells. The neurons subserving fine discrimina-
perature may be useful when the patient does not tol- tive touch are the largest cells in the DRG. Tactile
erate pinprick stimuli, has confusing or inconsistent sensation follows several different pathways within
responses to pain testing, or to help map an area of the central nervous system. The central processes
sensory loss. In some instances, the deficit is more enter the spinal cord via the medial division of the
consistent with temperature testing than with pin- posterior roots, and bifurcate into ascending and
prick. Temperature testing may not be very reliable descending fibers (Figure 32.4). Fibers carrying fine

Campbell_Chap32.indd 526 6/21/2012 5:03:47 PM


Nucleus ventralis
posterolateralis (VPL) of
thalamus

Gasserian
(trigeminal)
ganglion

FIGURE 32.4 The tactile pathways.

527

Campbell_Chap32.indd 527 6/21/2012 5:03:47 PM


528 SECTION F | THE SENSORY SYSTEM

discriminatory and localized tactile sensibility then, nucleus gracilis lie dorsolaterally level (homunculus
without synapsing, turn upward in the ipsilateral pos- in Trendelenberg). The lemniscal fibers are joined by
terior column. Fibers carrying crude touch synapse analogous fibers subserving facial sensation that have
within several segments of their point of entry, and decussated after synapsing in the trigeminal principal
the axons of the neurons of the next order cross to sensory nucleus in the pons. These fibers all termi-
the opposite ALS. Other tactile fibers have a synapse nate in the thalamus, from which the thalamocortical
in the posterior horn, and ascend in the dorsolateral radiations project to the somatosensory cortex. The
funiculus to the lateral cervical nucleus at C1-C2, distribution of the tactile impulses within the tha-
where axons of the next order neurons decussate and lamic nuclei and their radiation to the parietal cor-
join the ML. In the posterior columns, fibers from tex in general follow that for pain and temperature
the lumbosacral region aggregate near the midline, impulses.
and fibers from successively more rostral regions
aggregate in a progressively more lateral position,
producing somatotopic lamination, the reverse of
Clinical Examination
the STs (Figure 32.3). In the STs, the sacral fibers are There are many methods available for evaluating tac-
most lateral; in the posterior columns, the lowest fib- tile sensation. Light touch can be tested with a wisp
ers are most medial. All the fibers below about T8 are of cotton, tissue paper, a feather, a soft brush, light
grouped together in the fasciculus gracilis; analogous stroking of the hairs, or even using a very light touch
fibers above T8 form the fasciculus cuneatus. of the fingertip. Some appreciation of light touch
Anterolateral system fibers transmit light touch may be obtained by noting the responses to the blunt
and light pressure sensations, without accurate locali- end of the stimulus used to test pinprick.
zation. The posterior column fibers are concerned More detailed and quantitative evaluation can
with highly discriminatory and accurately localized be accomplished using Semmes-Weinstein filaments,
sensibility, including spatial and two-point discrimi- an asthesiometer, or von Frey hairs. These methods
nation. Because of the overlap and duplication of employ filaments of different thicknesses to deliver
function, and because of the multisynaptic path- stimuli of varying, graded intensity. For routine test-
ways for general tactile sensation, tactile sensibility ing, simple methods suffice. It is enough to determine
is the sensory modality least likely to be completely whether the patient recognizes and roughly localizes
abolished with lesions of the spinal cord, and distur- light touch stimuli and differentiates intensities. The
bances of it may fail to give localizing information. A stimulus should not be heavy enough to produce
myelopathy severe enough to abolish light touch will pressure on subcutaneous tissues. Ask the patient
often render the patient nonambulatory. to say “now” or “yes” on feeling the stimulus or to
Axons in the gracile and cuneate fasciculi syn- name or point to the area stimulated. Allowance must
apse with second-order neurons in the gracile and be made for the thicker skin on the palms and soles
cuneate nuclei at the cervicomedullary junction. The and the especially sensitive skin in the fossae. Similar
second-order neurons sweep anteriorly as internal stimuli are used for evaluating discriminatory sen-
arcuate fibers, cross the midline, and accumulate in sory functions such as tactile localization and two-
the ML. Within the medulla, the ML is a vertical point discrimination. It is best to avoid hairy skin
band of fibers situated along the median raphe; in the because the sensory stimulation due to hair motion
pons, the tract becomes more horizontal and shifts may be confused with the test stimulus; hairy skin is
to a ventral position; and in the mesencephalon, the exceptionally sensitive to touch. Two-point discrimi-
tract migrates to lie far laterally in an oblique posi- nation is considered both a delicate tactile modal-
tion. Somatotopic organization is maintained in the ity and a more complex sensation requiring cortical
ML. In the medulla, the fibers from the nucleus gra- interpretation.
cilis lie ventrally and those from the nucleus cunea- Using painless and noninvasive reflectance in
tus dorsally (homunculus erect). As the ML ascends vivo confocal microscopy of skin, investigators are
the brainstem, it moves from a vertical, paramidline able to visualize and quantitate Meissner’s corpuscles
position gradually to a horizontal position (homun- (MC) in dermal papillae. Comparing the density of
culus sits, then lies down). In the pons, fibers from MC may prove very useful for noninvasive detection
the nucleus gracilis lie laterally and those from cune- and monitoring of patients with sensory neuropa-
atus medially. In the midbrain, the fibers from the thy. Epidermal nerve fiber layer assessment on skin

Campbell_Chap32.indd 528 6/21/2012 5:03:51 PM


CHAPTER 32 | THE EXTEROCEPTIVE SENSATIONS 529

FIGURE 32.5 The Bumps device. Each square on the plate contains five colored circles; one randomly selected colored circle in
each square contains a bump. The 12 bumps have different heights. (Reprinted with permission from Kennedy WR, Selim MM, Brink
TS, et al. A new device to quantify tactile sensation in neuropathy. Neurology 2011;76:1642–1649.)

biopsy has been used to evaluate patients with small Cohen MS, Wall EJ, Brown RA, et al. 1990 AcroMed Award in
fiber neuropathies. Assessment of MC may bring basic science. Cauda equina anatomy. II: Extrathecal nerve roots
and dorsal root ganglia. Spine 1990;15:1248–1251.
such capability to the evaluation of large fiber neu- Cook AW, Nathan PW, Smith MC. Sensory consequences of com-
ropathies. Other detectable changes in neuropathy missural myelotomy. A challenge to traditional anatomical con-
include distortion of MC structure, focal thinning, cepts. Brain 1984;107:547–568.
Defrin R, Ohry A, Blumen N, et al. Sensory determinants of ther-
or loss of myelin and short myelin internodes.
mal pain. Brain 2002;125:501–510.
Using the finger pad, detection of small bumps Dyck PJ. Enumerating Meissner corpuscles: future gold stan-
on a smooth surface is mediated by MC and large, dard of large fiber sensorimotor polyneuropathy? Neurology
myelinated nerve fibers. A simple device called 2007;69(23):2116–2118.
Dyck PJ, O’Brien PC, Bushek W, et al. Clinical vs. quantita-
Bumps appears to be a rapid, sensitive, and inexpen- tive evaluation of cutaneous sensation. Arch Neurol 1976;
sive method to quantitate tactile sensitivity of the 33:659.
finger pads (Figure 32.5). Patients with neuropathy Dyck PJ, Zimmerman IR, O’Brien PC, et al. Introduction of
had lower MC density on skin biopsy and elevated automated systems to evaluate touch-pressure, vibration, and
thermal cutaneous sensation in man. Ann Neurol 1978;4:
threshold for detecting bumps compared to controls. 502.
Friehs GM, Schrottner O, Pendl G. Evidence for segregated pain
and temperature conduction within the spinothalamic tract.
J Neurosurg 1995;83:8–12.
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