‫‪.

Biomechanics‬‬ ‫‪of nerves‬‬

‫اسم الطالب ‪ :‬محمد عماد حمدي سيد‬ ‫‪‬‬
‫الفرقة ‪ :‬الثانية‬ ‫‪‬‬
‫‪ID :211080103‬‬ ‫‪‬‬
 :Introduction

There has been an emergence in physical therapy of evaluation and intervention based
on neurodynamics, the relationship between nerve physiology and nerve
mechanics.1 To advance the clinical care of people with nerve injuries, it is essential
to understand peripheral nerve structure and plasticity. The purpose of this appraisal is
to review the structural and biomechanical properties of peripheral nerves and then to
discuss how nerves respond to physical stresses. We will expand on the Physical
Stress Theory presented by Mueller and Maluf2 and discuss the structural and
biomechanical modifications seen in nerves exposed to various levels of physical
stress. We hold the premise that physical therapists who understand the adaptive
responses of nerves to specific physical stresses will be better prepared to provide
reasoned interventions to mo dify specific aspects of the stresses. These
knowledgeable therapists also may educate patients in injury prevention and self-care
.and thus significantly improve function and quality of life
 :Anatomy of nerve
For the most part, nerves in the body are relatively unprotected by any bony structures
in comparison to the brain and spinal cord that have the mechanical protection by the
skull and spinal column, respectively. As such, nerves are an organized collection of
axons with connective tissue that provides compressive and tensile strength (Fig. 1).
The axons in peripheral nerves are organized into fascicles, or bundles that
collectively make up a nerve. At each organizational level, there is a layer of
connective tissue (Fig. 1). The endoneurium envelopes each axon, the perineurium
encloses each fascicle, and the epineurium forms a tough, fibrous sheath around the
.entire structure of the nerve

Other cells, such as Schwann cells support the chemical and homeostatic environment
of the neurons to facilitate axonal survivability and electrical conduction along the
axons. While those support cells provide an environment that is conducive to and
maintains the normal function of axons, they provide minimal mechanical
contributions for the structural integrity of axons. As such, they will not be discussed
in this chapter. Within each nerve, the fascicular bundles and the axons course
through the nerve in an undulating manner; this undulation allows for some
elongation along the nerve without imposing significant loading along the lengths of
the fascicles and axons as they may become straightened when the overall nerve is
pulled in tension [75, 80]. The structural organization and mechanical properties of
the connective tissue of the nerve are able to preserve the integrity of the relatively
fragile axons in the nerves under normal physiologic conditions such as joint motions
.and muscle elongation

Together, the epineurium, perineurium, and endoneurium provide mechanical

protection when tension and/or compression are applied to the overall nerve structure.
Type I and Type III collagen, as well as elastic fibers, comprise the connective tissue
of the epineurium [31, 80]. The dense network of collagen and elastic fibers that make
up epineurium tissue can absorb shock by dissipating the compressive forces and
protecting nerves from such compression to the nerve structure as a whole [75]. While
it has been hypothesized that the epineurium does provide some protection to the
nerve in tension, it does not appear to be the primary contributor to the tensile
mechanical strength of the nerve [23, 66]. In all but the most-peripheral of nerves, the
nerve contains more than one fascicle [71]. For nerves with more than one fascicle,
the epineurium is divided into the epifascicular epineurium, surrounding the entire
nerve, and the interfascicular epineurium that separates the individual nerve fascicles.
The perineurium surrounding each fascicle is the primary contributor to the nerve’s
tensile strength and elasticity [66, 75]. This layer contains Type I and Type II collagen
fibers, as well as elastic fibers oriented in circumferential, oblique, and longitudinal
orientations, which together can potentially provide multidirectional tensile strength
to the fascicles [31, 80]. The endoneurium that surrounds each axon is composed of
collagen (Types I, II, IV) and provides a nominal degree tensile strength [75, 79, 80].
The three layers of connective tissue that surround nerve tissue act in concert across
scales and directions to provide the compressive and tensile strength collectively
necessary to protect the axons they surround during normal motions and physiologic
.loading scenarios

.
:Biomechanical properties of nerve

Under normal physiological conditionsimposed by posture and movement,nerves are

exposed to various mechan-ical stresses. Stress is defined as forcedivided by the area
over which itacts9,23–25and can be applied to a nerveas tensile, compressive, or
shear stressor as a combination of stresses (Fig. 4).Tensile stress may be applied to
tissueseither parallel or perpendicular to thelength of the nerve, causing
respectivelongitudinal or transverse stress in thenerve. When joint motion causes
elon-gation of the nerve bed, the nerve isinherently placed under tensile stressand
accommodates the stress by bothelongating and gliding.15The deforma-tion or change
in nerve length inducedby longitudinal tensile stress is calledstrain and is expressed
typically as per-cent elongation.23,26–28Displacementor gliding of a nerve relative to
thesurrounding nerve bed is called excur-sion.29–31The direction of excursion may
be longitudinal or transverse, or both, relative to thenerve tract31,32and is measured
in millimeters.The direction and magnitude of nerve excursion aredependent upon the
anatomical relationship betweenthe nerve and the axis of rotation in the
movingjoint.29,33When the nerve bed is elongated, the nerve isplaced under
increased tensile stress. With the elonga-tion of the nerve bed, the nerve glides toward
themoving joint,1,33,34a movement termed convergence.1Conversely, if the nerve
bed tension is relieved duringjoint motion, the nerve will realign along the
shortenednerve bed by gliding away from the moving joint, amovement termed
divergence.33Convergence in themedian nerve may be demonstrated during elbow
exten-sion (Fig. 5). The motion elongates the bed of themedian nerve, causing the
nerve segment in the arm toglide distally toward the elbow and the nerve segmentin
the forearm to glide proxi mally toward the elbow. Incontrast, elbow extension
relieves the tensile stresses inthe ulnar nerve bed, causing the ulnar nerve to
divergeaway from the elbow (Fig. 5). With limb movement,nerve excursion occurs
first in the nerve segment imme-diately adjacent to the moving joint. As limb
movementcontinues, excursion occurs at nerve segments that areprogressively more
distant from the moving joint.29,33Similarly, the magnitude of excursion is greatest
in thenerve segments adjacent to the moving joint and is leastin the nerve segments
distant from the joint(Fig. 5).29,33,34The magnitude and direction of medianand
ulnar nerve excursion for motions of the joints ofthe upper limb commonly used in
clinical assessment areshown in Table 1.As tensile stresses cause measurable nerve
excursion,they simultaneously produce changes in strain within thenerve. Elongation

of a nerve bed during joint movementwill cause an increase in nerve strain (Fig. 6).
Themagnitude of increased strain with limb movement isgreatest in the nerve segment
closest to the moving joint(Fig. 6).33–35The mechanical behavior of a nerve seg-
ment under longitudinal tensile force may be describedby a load-elongation
curve,36or by a stress-strain curve ifthe force is divided by the cross-sectional area of
thenerve, and elongation is expressed as a percentage of thestarting length (Fig. 7).
When a load is first applied to aresting nerve, the nerve lengthens markedly relative
tothe applied load, as shown in the “toe region” of thecurve. Structurally, the minimal
longitudinal tensile loadresults in straightening of the wavy connective tissue
andaxons in the endoneurial compartment and in thedisappearance of the periodic
light-reflecting bands inthe epineurium.12,37,38As the tensile load is increased,the
nerve elongates at a steady rate, as demonstrated bythe linear region of the load-
elongation curve (Fig. 7).The slope of this portion of the curve is a measure of the
resistance of the nerve to deformation and is termedstiffness in the load-elongation
curve or modulus of elastic-ity in the stress-strain curve.A steep slope indicates
thatthe nerve has more stiffness, has less elasticity, and is lesscompliant than a nerve
with a shallower slope. At acertain point, the amount of applied load starts
topermanently deform particular elements of the nerve.This ultimate stress or ultimate
strain represents thetransition between the recoverable (elastic) strain andplastic
(permanent) deformation areas of the load-elongation curve. Finally, in the plastic
region of thecurve, the nerve reaches its ultimate elongation andundergoes mechanical
failure. Minor increases in tensileload create significant elongation of the nerve
becauseof the failure of the infrastructure of the nerve, includ-ing perineurial
components. There are fewer intactstructural elements to provide resistance, and at
thispoint, the nerve behaves like a viscous material.36A nerve in situ is under some
tensile load, as evidencedby the fact that a nerve in situ retracts when severed.
Thepercent change in length is termed the in situ strain,37which corresponds
approximately to the transitionbetween the toe region and the linear region of
thestress-strain curve. The magnitude of the in situ strain isdependent upon the
configuration of the nerve bed. Ina slackened position, such as in the rabbit tibial

nervewhen the knee and ankle are each maintained at 90degrees of flexion, the in situ
strain is 11%.37Extensionof the knee or dorsiflexion of the ankle places greater
tensile force on the nerve in the elongated nerve bed,and the in situ strain increases
from the original 11%.Interestingly, it has been suggested that the toe region ofthe
stress-strain curve may be a property of excisednerves, as in situ nerves immediately
enter the linearportion of the stress-strain curve when placed underincreasing tensile
stress from a “rest” position.39Becausethe in situ strain is a direct reflection of
cumulative nervepositioning across multiple joints, one must consider theeffect of
trunk, neck, and limb positioning during clin-ical assessment and intervention to
minimize physicalstress on an injured nerve.To elongate a nerve, thus increasing its
strain, the tensilestrength inherent in the nerve from elastic and connec-tive tissues
must be overcome. Elongation of a nerve isknown to cause a reduction in the cross-
sectional area, aproperty called transverse contraction15(Fig. 4). This prop-erty results
in increased pressure in the endoneurialcompartment. A recently proposed theoretical
modelsuggested that the outer connective tissue tube or sheathconstraining the inner
pressurized neural core contrib-utes significantly to the biomechanical properties of
anerve placed under tensile strain.15,39Upon elonga-tion of a nerve, the increased
pressure produced in theneural core will resist the transverse contraction and will
contribute to the stiffness of the nervewhen stretched.15When the tensilestress is
removed, it is likely that acombination of elasticity of the connec-tive tissues and
pressure within the neu-ral core will allow recoiling of the nerveto nearly the original
cross-sectionalarea and length. Recent studies40,41defined the core-sheath interface
as theinnermost cell layer of the perineuriumand suggested that the interface pro-vides
some minimal resistance to elon-gation. With increasing tensile load,structural
separation occurs first in thecore-sheath interface, then in the axonsand connective
tissues in the endoneur-ial core, and finally in the cells andconnective tissues of the
perineurialand epineurial sheath. It is importantto understand that diffuse damage
toaxons in the endoneurial core mayoccur long before visible damage to
theepineurium.There are a number of factors thataffect nerve compliance and thus
dic-tate the level of strain, excursion, andtransverse contraction in the nerve dur-ing
limb movement.12,24,42First, arecent study43measured greater nervecompliance in
nerve segments thatcross joints than in segments that donot cross joints. The median
and sciaticnerves exhibit more strain in situ andless stiffness ex vivo in the
segmentsthat cross the elbow and hip than indistal segments that do not cross

therespective joint. Although the bio-mechanical findings correlated withfascicle

number and cross-sectionalarea of extrafascicular connective tis-sues in the sciatic
nerve, there was nosuch correlation in the median nerve.It was initially thought that
there aremore fascicles and connective tissues where nerves crossjoints12and greater
stiffness in nerve segments withmultiple fascicles.9,12However, these notions do
notseem to hold true for all nerves at all joints.43Thus,internal neural structure is but
one factor affecting nervecompliance.Second, nerve stiffness is greater in long nerve
sectionsand in nerve sections with numerous branches.15Sever-ing nerve branches or
vessels but leaving the nerve insitu results in increased compliance and decreased
stiff-ness.15Excising the same nerve completely from its nerve bed results in further
increases incompliance, likely because of reducedfriction between the nerve and
paran-eural tissues15and possibly because of aloss of internal pressure.39Third, nerve
stiffness is greater when anerve is elongated rapidly rather thanslowly. In addition, the
ultimate strainat the point of failure appears to bedependent on the rate of
elongation.Haftek36measured compliance inexcised rabbit tibial nerves elongated
at0.5 mm per minute to the point offailure. At this point, the nerves had amean
ultimate elongation, or strain, of55.7%. Rydevick et al10elongatedexcised rabbit tibial
nerves at 1.0 cmper minute, or 20 times faster, andfound a mean ultimate strain of
38.5%.The reduction in the ultimate strainsuggests that nerves elongated at agreater
rate exhibit a reduction in theirability to tolerate elongation. Theserate-dependent
effects are characteris-tic of tissues that exhibit viscoelasticbehavior.When a nerve is
placed under tensionand maintained at that new fixedlength over time, there is a
reduction inthe tension in the nerve or the forcerequired to maintain the fixed

length.The observed reduction in tension maybe plotted in a stress-relaxation

curve(Fig. 8).25,44The majority of relaxationoccurs in the first 20 minutes of
fixedelongation.25,44Stress relaxation innerves that are stretched slowly is
greaterthan in nerves that are stretchedrapidly.25,37,44– 46This phenomenon
wasobserved when comparisons were madefor rabbit tibial nerves stretched at differ-
ent rates to lengths 6% longer than theirresting lengths. Over the 60-minuterelaxation
time, there was a 57% reduc-tion in stress in nerves elongated at0.08% per
second,45but only a 34%reduction in stress in nerves elongated at3.0% per
second.44The same effect wasnoted for nerves that were subjected to12%
strain.44,45A phenomenon analo-gous to stress-relaxation behavior, com-monly
referred to as “creep,” is seenwhen a nerve is maintained under a fixedtensile load.
Nerve tissue elongates gradually under these loading conditions. Both stress
relaxationand creep are used to quantitatively describe the viscoelas-tic behavior of a
material46and may provide some protec-tion for nerves during postures in which
nerves are underprolonged lengthening and tensile loads.Tensile stress applied
repetitively also may alter thestress-strain curve. At strains below 8%, repetitive
stretchhas no effect on the stress-strain curve, as shown in arabbit model of sciatic
nerve strain in situ.47However, anerve stretched repetitively to 8% or 10% strain
exhibitsa reduced slope of the stress-strain curve, indicating thatthat nerve undergoes
less stress with successive elonga-tions because of increased compliance and
decreasedstiffness. Note that repetitive application of a consistenttensile stress will
result in a progressive increase in nervestrain. As discussed in the section on the
responses ofnerves to physical stress, high levels of strain will result inphysiological
and structural alterations in the nerve. Thisinformation provides a rationale for
incorporating active it's modification into patient education in addition to tensile
stress, nerves are exposed staticallyand dynamically to compressive stresses. As
mentionedpreviously, the laws of physics dictate that the cross-sectional area of a
cylindrical object is reduced as thecylinder is elongated. As a nerve is elongated
undertensile force, the nerve undergoes transverse contrac-tion, which is resisted by
the fluid and nerve tissuecontained within the connective tissue
sheath.15,39Themagnitude of the transverse contraction stress is greatestat the center
of the elongating segment15(Fig. 4).Nerves also may be compressed externally by
approxi-mation to adjacent tissues, such as muscle, tendon, orbone, or by pressure
increases in the extraneural envi-ronment. Compression of a nerve segment causes
dis-placement of its internal contents in transverse andlongitudinal directions. As
shown in rat nerve, extra-neural compression causes an immediate displacementof
endoneurial fluid to the edges of a compressive cuffover 5 to 10 minutes and a much
slower displacement ofaxonal cytoplasm over the course of hours.48The dam-age to
axons and myelin is greatest at the edges of thecompressed zone,48,49where the shear
forces are heights 50In a carefully controlled study,Dyck and
colleagues48demonstratedthat under a compressive cuff, length-ening of internodes,
cleavage of para-nodal myelin, and myelin laminae over-lapping nodes occurred. At
the edgesof the cuff, however, myelin retractionwith resultant widening of nodes
andparanodal demyelination occurred.These structural alterations in myelinmay be
expected to result minimally inimpaired impulse conduction or maxi-mally in
demyelination and a conduc-tion block.In response to biomechanical stressesplaced
on a nerve as an individualassumes a posture or movement, thenerve follows the path
of least resis-tance.29Combinations of tensile, shear,and compressive stresses result in
com-binations of nerve excursion, strain, andtransverse contraction. Because the bio-
mechanical forces on the nerve are sointricately linked, the sequencing andrange of
joint movement affect themagnitude and direction of excur-sion,27,29the magnitude
of nervestrain,27,29,35and the degree of trans-verse contraction at different sitesalong
the nerve.27An extensive reviewof the literature has allowed us to for-mulate tables
of the normal ranges ofexcursion (Tab. 1) and strain (Tab. 2)in 2 upper-extremity
nerves measuredsubsequent to movements of the upperextremity. It is important to
note thatthe magnitude of strain achieved withnormal range of motion approaches
orexceeds the magnitude known to resultin physiological changes in the nerve.In
comparing data from different stud-ies, one must make note of the testposition,
location of the measurement,measurement tools, and type of speci-men used in each
study.Simultaneous nerve excursion, strain,and transverse contraction may be seenin
the ulnar nerve as an example ofresponses to physical stresses imposedduring
movements of the upper limb.When the upper limb is maintained ina position of 90
degrees of shoulderabduction and 90 degrees of shoulderexternal rotation with the
wrist neutral and when the elbow is moved from 90degrees of flexion to full
extension, theulnar nerve bed is shortened and thetensile stress on the nerve is
decreased.With this motion, there is divergence ofthe ulnar nerve away from the
elbow(Fig. 5), decreased nerve strain, espe-cially at the elbow (Fig. 6), anddecreased
compression within the cubi-tal tunnel.27,29,33When the wrist then isextended from
neutral to full exten-sion, the ulnar nerve bed is lengthened,resulting in convergence
of the nervetoward the wrist (Fig. 5), an increase innerve strain (Fig. 6), and
transversecontraction greatest in the nerve seg-ment across the carpal bones and at
thetunnel of Guyon.27,29,33The magnitudeof nerve strain and excursion will
begreatest near the wrist, and the fascicleswill rearrange as the nerve assumes
aflattened oval shape. Because the nervedoes not lie directly on the rotationalaxis of
joint motion, the fascicles far-thest from the axis will undergo greaterstrain than those
closer to the center ofrotation51The magnitude of these bio-mechanical effects will
depend uponthe rate at which the limbs are moved,the time spent in each position, and
thetemporal aspects of the movement ifthe motion is repetitive. A
thoroughunderstanding of the anatomical andbiomechanical relationships of
nervesand surrounding tissues is necessary tointerpret the responses of patients
.topostures and movements of the trunk,head, and limbs