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THE NEUROLOGY OF JOINTS
Arris and Gale Lecture delivered at the Royal College of Surgeons of England
on
17th February 1966
by
Barry Wyke, M.D., B.S.
Neurological Laboratory, Department of Applied Physiology,
Royal College of Surgeons of England
'But the expression of a well-made man appears not only in his face,
It is in his limbs andjoints also, it is curiously in thejoints of his hips and wrists,
It is in his walk, the carriage ofhis neck, theflex of his waist and knees....."
-WALT WHITMAN, Leaves of Grass (1891-2)
THE THEME I propose to pursue in this lecture-the neurology of joints-
involves consideration of the innervation of the joints of the skeleton, and
a description of the role of this innervation in the regulation of muscular

Reprodu.ced by permission of the Worshipful Company of Barbers

Fig. 1. A portrait by Richard Greenbury (now in the Guildhall Art Museum,
London) of Mr. Edward Arris (on the right) in attendance at one of the Muscular
Lectures endowed by him. The lecture was delivered at Surgeon's Hall by Sir
Charles Scarborough (on the left) in 1649-although the picture was not painted
until 1651.
activity. This would seem an appropriate modern counterpart to the
three-hundred-year-old intentions of the co-founders of the Arris and
Gale trusts (Fig. 1) (Stack, 1963)-which were intended at their inception
to provide a series of lectures on muscular and osteological topics
(Copeman, 1963; Dobson, 1964).
25
 BARRY WYKE
Our interest in the subject of articular neurology arises from realization
that the neurology of joints, in spite of its obvious relevance to several
branches of surgery, and to physical medicine generally, is a field of
knowledge that has never been adequately explored*; and for this reason,
when the new Neurological Laboratory in this College was opened three

(a)

(c)

(b)
Fig. 2. Dissections of the specific articular nerves supplying the knee joint (in the
cat). (a) The left posterior articular nerve (indicated by arrow), arising from the
posterior tibial nerve (n) at the lower end of the popliteal fossa, and branching to
enter the posterior capsule of the joint (c) on the right. x 1. (b) The medial
articular nerve exposed on the antero-medial aspect of the right thigh after removal
of the sartorius muscle. In this instance, the nerve arises from the obturator nerve
trunk and passes anteriorly through the adductor musculature to the front of the
thigh. It branches just above the medial condyle of the femur below, close to the
medial superior genicular artery (a): (p) is the patella. x i. (c) The lateral articu-
lar nerve (indicated by large arrow) supplying the left knee joint. It arises from the
lateral popliteal nerve (n) on the surface of the lateral head of the gastrocnemius
muscle (G) and ascends to branch into the inferior edge of th- lateral capsule of the
joint (c). A second articular branch (indicated by small arrow) arises distally from
the lateral popliteal nerve, to supply the superior tibio-fibular joint (t). x .
years ago, we decided to devote its excellent resources to a co-ordinated
attack on the subject. To this end, we have pursued the general topic
of articular neurology along three parallel lines, employing similar methods
of investigation in each case:
* Thus, in a recent comprehensive monograph on joints, Barnett et al.
(1961) were
forced to write: " In the present state of knowledge, it is impossible to define with any
precision the role of the receptors in the joint tissues in the control of posture, movement,
tone of muscles and so forth."
26
 THE NEUROLOGY OF JOINTS
1. Studies of the limb and vertebral joints, whose neurological func-
tions are important to orthopaedic and neurological surgeons, and to
practitioners of physical medicine.
2. Studies of the laryngeal joints, and of the ossicular joints of the
middle ear, in relation to the mechanisms of speech and hearing-a subject
that is of obvious significance to otolaryngologists and speech therapists.
3. Studies of the temporo-mandibular joints, whose neurology is rele-
vant to many problems of oral and facio-maxillary surgery.
We are fortunate in having made a certain amount of progress in each
of these fields, the details of which are recounted in the papers and mono-
graphs listed at the end of this lecture. In the course of this work,
however, it has become clear that the neurological aspects of each of these
systems of joints share certain basic features in common; and it is to these
more general aspects of articular neurology that I here wish to draw
attention.
THE INNERVATION OF JOINTS
As with every neurological investigation, the first step in this field is
anatomical, and involves the definition of the structural features of the
nerve supply of each system of joints.
Extrinsic innervation
The extrinsic innervation of each joint is being investigated by macro-
and micro-dissection in animals and man. In this way, it has been found
that each joint has a dual pattern of nerve supply-first, by specific
articular nerves that reach the joint capsule as independent branches of
adjacent peripheral nerves, often (but not exclusively) in company with the
articular blood vessels; and secondly, by non-specific articular branches of
related muscle nerves. These latter arise within some of the muscles
that are attached to each joint capsule as intramuscular branches of various
muscle nerves, and reach the joint by running through the substance of the
muscles, embedded in the interfascicular connective tissue. Two examples
from different joint systems will serve to illustrate these points.
1. The knee joint is supplied by three specific articular nerves-a
posterior, a medial and a lateral articular nerve, derived respectively from
the posterior tibial, femoral (or obturator) and lateral popliteal nerves.
Dissections of each of these are displayed in Figure 2-the posterior
articular nerve (in the popliteal fossa) on the upper left (Fig. 2a), the medial
articular nerve (on the antero-medial aspects of the thigh) on the right
(Fig. 2b), and the lateral articular nerve (on the supero-lateral aspect of the
leg) on the lower left (Fig. 2c). The knee joint is also supplied by numerous
non-specific intra-muscular branches derived from the nerves to the
sartorius and quadriceps musculature, and by a few twigs derived from the
cutaneous nerves innervating the skin over the joint.
2. Again, the laryngeal joints are each supplied by specific articular
branches of the recurrent and internal laryngeal nerves, and by intra-
27
 BARRY WYKE
muscular twigs running within some of the muscles attached to the laryn-
geal cartilages. Thus, Figure 3 shows an articular branch of the internal
laryngeal nerve that innervates the crico-arytenoid joint.
As similar relationships appear to obtain with all other joints, it seems
that the basic implications of Hilton's Law (1863)* are applicable to all the
joints of the body, and not merely to those of the limbs-although modi-
fications of detail, which I cannot describe here, have to be made in apply-
ing this law to the individual joints.

Fig. 3. The innervation of the crico-arytenoid joint (cat). The articular nerve
(indicated by arrow) arises from the internal laryngeal nerve (I). The larynx is
viewed from the left, after removal of the thyroid cartilage (the cut edges of which
are seen at TC). (T) is the trachea, with the recurrent laryngeal nerve (R) beside it.
x2

Constitution of articular nerves

Neurohistological studies have shown that each articular nerve contains
a mixture of myelinated and unmyelinated nerve fibres, whose diameters
range from less than lI up to 17ji. Considered in combination with the
results of oscillographic analyses of the impulse traffic in the articular
nerves, of electrical stimulation procedures, and of other neurophysio-
* Hilton stated: " The same trunks of nerves whose branches supply the groups of
muscles moving a joint, also furnish a distribution of nerves to the skin over the in-
sertions of the same muscles and-what at the moment more especially merits our
attention-the interior of the joint receives its nerves from the same source."
28
 THE NEUROLOGY OF JOINTS
logical investigations to be described later, these studies suggest that the
fibres in articular nerves generally may be subdivided into three principal
size categories, as indicated in Table I, each of which has specific functional
correlates.
First, a large proportion (at least 45 per cent) of the total number of
fibres in each articular nerve has diameters of less than 51. Most of these
small myelinated and unmyelinated fibres, which are embraced in Group
IlI of the Table, are afferent in function and subserve articular pain
sensation; but a small proportion of the unmyelinated fibres in this group
consists of visceral efferent fibres of sympathetic origin, that innervate the
articular blood vessels that is, these latter are articular vasomotor nerve
fibres. Thus far, we have no anatomical or physiological evidence of the
presence of secretomotor fibres in the articular nerves-or, indeed, of any
direct nervous influence on the production of synovial fluid.
TABLE I
GENERAL COMPOSITION OF ARTICULAR NERVES
Group Diamleter Structure Function
number rai;ge
(p)
I 13-i7 Large myelinated Mechanoreceptor afferent
(from o:nt ligaments)
II 6- 12 Medium myelinated Mechanore eptor afferent
(from fibrous capsule and fat
pat ))
III 2-5 Small myelinated Pai:t afferent
<2 Unmyelinated Pain afferent
Vasomotor efferent
Secondly, another large proportion (some 45 to 55 per cent of the total)
consists of medium-sized myelinated fibres between 6p and 12i in diameter,
and forms Group II of our classificat on. All of these fibres are mechano-
receptor afferents, innervating small corpuscular endorgans located in the
fibrous capsules and fat pads of the joints that will be described in a
moment.
Thirdly, a small proportion (some 10 per cent, or less) consists of large
myelinated fibres between 13 and 17p in diameter, and forms Group I
of our classification. These also are mechanoreceptor afferents, inner-
vating large corpuscular endorgans that are confined to the joint ligaments.
This last group of fibres is absent from the articular nerves of the larynx
and middle ear, but is represented in most of the other articular nerves that
we have examined.
In the case of the limb and vertebral joints, the nerve cell bodies from
which the articular afferent fibres derive are located in the segmentally
related dorsal root ganglia; and in the case of the laryngeal joints, they are
situated in the ganglion nodosum of the vagus nerve. We have not
yet established the location of the nerve cell bodies of the afferent
fibres innervating the temporo-mandibular and ossicular joints-but
29
 BARRY WYKE
we have a priori reasons for expecting that they will be found in the
mesencephalic nucleus of the fifth nerve in the case of the mechanoreceptor
afferents from the temporo-mandibular joints, and in the trigeminal
ganglion in the case of the pain afferent fibres from these same joints.
Articular receptor endings
Using specially modified neurohistological staining techniques* in-
volving gold, silver and methylene blue procedures, it has been possible to
show that the articular nerve fibres just described terminate in four
structurally distinct varieties of ending within the articular tissues (Fig. 4).
Correlated neurophysiological investigations, by ourselves and by others,
have further revealed that these morphologically distinct categories of
nerve ending possess individually distinct behavioural characteristics;
and, on this basis, we have proposed a general structure-activity scheme
for the classification of articular nerve endings, the basic criteria for which
are summarized in Table 1I.
Type I receptors
The Type I receptors, a group of which is shown in Figure 4a, are
globular or ovoid corpuscles, similar to those described originally by
Ruffini (1905) in subcutaneous and fascial connective tissues. They are
numerous in relation to all the limb joints, the apophyseal joints of the
vertebral column, and the temporo-mandibular joints; but in the larynx,
they are restricted to the crico-arytenoid joint, being absent from the
thyro-epiglottic, thyrohyoid and cricothyroid joints. In the middle ear,
we have likewise failed to find them in the ossicular joints-but this nega-
tive observation cannot yet be considered definitive, as our histological
study of this system of joints is not yet complete. In the limbs, the Type I
receptors appear to be more numerous in the proximal (for example, in
the shoulder and hip) joints than in the distal joints; and in the spine,
they appear to be more numerous in the joints of the cervical region than
elsewhere-but again, this distribution requires further statistical analysis
before it can be regarded as settled.
In each joint in which they are present, the Type I receptors are located
mainly in the superficial (that is, in the external) layers of the fibrous cap-
sule (as shown in Fig. 4a), within which they are distributed tridimension-
ally in clusters of up to six corpuscles. Each member of such a cluster
is supplied by terminal branches from a single Group II myelinated parent
afferent axon some 6-9pi in diameter, that enters the joint capsule in one
of the articular nerves-as can be seen in Figure 4a. Within the capsule
of each individual joint, the clusters of Type I receptors show regional
differences in distribution density, in that they are more numerous on
* We are deeply grateful to Dr. M. J. F. Fitzgerald (Department of Anatomy,
University College, Cork, Eire) and to Dr. C. Coers (Departement de Neurologie,
Universit6 Libre de Bruxelles) for advice and assistance in the development of some of
these procedures for use on articular tissues.
30
 THE NEUROLOGY OF JOINTS

Fig. 4. The four varieties of articular receptor nerve ending (descriptions in text),
as seen in the articular tissues of the cat. (a) A Type I receptor cluster (one
corpuscle in which is indicated by arrow), embedded in the superficial surface of
the fibrous capsule of the joint. The myelinated afferent nerve fibre (N) can be
seen joining an articular nerve trunk (A) on the right. Wyke gold chloride method.
x 350. (b) A Type II corpuscle (indicated by arrow) in the joint capsule, at the
junction between the fibrous capsule (FC) and the sub-synovial fibro-adipose
tissue (FA). The myelinated afferent nerve fibre (N) can be seen joining an
articular nerve branch at (A). Fitzgerald frozen silver method. x 150. (c) A
Type III receptor corpuscle (indicated by arrow) on the surface of a collateral
ligament (L), near its fusion with the fibrous capsule of the joint (FC). (N)indicates
the myelinated afferent nerve fibre. Fitzgerald frozen silver method. x150.
(d) Portion of the Type IV plexus system in the fibrous joint capsule, the super-
ficial (external) surface of which is seen above and to the left. The fine nerve
fibres, weaving amongst the small blood vessels, arise from myelinated and unmye-
linated parent nerve fibres within the fibrous capsule. Wyke gold chloride method.
x 150. (e) Type IV free nerve endings ramifying amongst the fibres of a joint
ligament. Fitzgerald frozen silver method. x 150.
31
 BARRY WYKE
those aspects of the joint capsule that undergo the greater changes in
stress during natural joint movement.
Physiologically, the Type I corpuscles behave as low-threshold, slowly
adapting mechanoreceptors responding to the mechanical stresses ob-
taining in the part of the fibrous capsule in which they lie. For this reason,
a proportion of the Type I receptors in each joint capsule is always active
in every position of the joint, even when it is immobile. This resting
discharge usually has a frequency of some 10-20 impulses per sec., and is
generated partly by the stresses created regionally within the joint
capsule by the varying degrees of tone in the muscles attached to it, and
partly by the overall capsular stress created by the fact that the intra-
capsular (i.e. the intra-articular) pressure is normally some 5-10 mm. Hg
less than the external atmospheric pressure. Alterations-either increases
TABLE II
CLASSIFICATION OF ARTICULAR RECEPTOR SYSTEMS
Type Morphology Location Parent nervefibres Behavioural characteristics
I Thinly encapsulated globular Fibrous capsule Small mvelinated Static and dynamic
corpuscles (I 00 x 40p), in of joint (mainly (6-9u) mechanoreceptors; low-
clusters of 3-6 corpuscles superficial layers) threshold, slowly adapting
JI Thickly encapsulated conical Fibrous capsule Medium myelinated Dynamic mechanoreceptors
corpuscles (280u x 120Au), of joint (mainly (9-12A) low-threshold, rapidly
in clusters of 2-4 corpuscles deeper layers). adapting
Articular fat pads
III Thinly encapsulated fusiform Joint ligaments Large myelinated Dynamic mechanoreceptors-
corpuscles (600g x 100l) (intrinsic and (13-17,) high-threshold, very slowly
extrinsic) adapting
IV Plexuses and free nerve Fibrous capsule. Very small myelinated Pain receptors; high-
endings Articular fat pads. (2-5A) threshold, non-adapting
Ligaments. Walls Unmyelinated
of blood vessels (<2,u)
or decreases-in the rate of this resting discharge occur whenever the joint
is moved actively or passively, whenever the tone in the related muscles
changes isotonically or isometrically, or whenever the pressure gradient
between the interior of the joint and the atmosphere is altered sufficiently.
When such changes in capsular stress are imposed, the Type I receptors in
the affected regions of the joint capsule respond with an abrupt rise or fall
in their resting discharge rate, after which they adapt slowly (over
several seconds) to a new discharge rate that is determined by the new
degree of stress now prevailing in the affected parts of the joint capsule.
In the light of these data, then, the Type I articular receptors can be cate-
gorized as static and dynamic mechanoreceptors, whose discharge pattern
signals static joint position, intra-articular pressure changes, and the
direction, amplitude and velocity of joint movements.
Type II receptors
The Type II receptors, one of which is illustrated in Fig. 4b, are elon-
gated conical corpuscles, with a thick, multi-laminated connective tissue
capsule enclosing a single (or sometimes multistranded) unmyelinated
nerve terminal that ends in a bulb or Y-shaped bifurcation near the apex
32
 THE NEUROLOGY OF JOINTS
of the corpuscle. * Within the capsule of the endorgan, the terminal axon
is ensheathed by a single palisade-like layer of clear, columnar cells; and
it is derived from an extra-corpuscular myelinated axon that is about 5j
in diameter. These corpuscles have been observed in all the joints we
have examined, without exception. In the joints of the larynx and middle
ear they constitute the sole corpuscular endorgan present (except for the
few additional Type I corpuscles mentioned earlier as being present in
the capsule of the crico-arytenoid joint); and in the limbs, they are rela-
tively more numerous in distal joints (for example, in the ankle joint) than
in more proximal joints. They are also particularly numerous in the
temporo-mandibular joints.
Like the Type I receptors, the Type II corpuscles are found in the
fibrous capsules of all the joints, partly in the superficial layers-where,
in the case of the limb, vertebral, temporo-mandibular and crico-arytenoid
joints, they are mixed with varying numbers of Type I corpuscles. Most
of the Type II corpuscles are located, however, in the deeper (that is, in the
internal) layers of the fibrous capsules of the joints, particularly (as can be
seen in Fig. 4b) at the border between the fibrous capsule and the sub-
synovial fibro-adipose tissue, where they often lie alongside, or coil around,
the articular blood vessels. They are distributed in each joint capsule in
clusters usually of 2-4 corpuscles, each member of such a cluster being
innervated (as shown in Fig. 4b) by a branch of a Group II parent mye-
linated articular nerve fibre whose diameter is between 9p and 12p. Similar
clusters of Type IL endings are present also on the surfaces of all the fat
pads related to the limb and temporo-mandibular joints, whether these be
intra-articular or extra-articular.
The Type II corpuscles behave as low-threshold, rapidly adapting
mechanoreceptors. They are entirely inactive in immobile joints,
becoming active only at the onset or cessation of joint movement-that is,
at the moment at which sudden changes of stress occur in the regions of
joint capsule or fat pad in which they lie. When they are so stimulated,
each cluster of Type II receptors emits a brief, high-frequency burst of
impulses into the related afferent axon that lasts less than one second;
and in rapidly moving joints, such as those in the larynx, the duration of
this impulse burst may be less than 0.5 sec. Furthermore, as the diameter
of the afferent nerve fibres innervating the clusters of Type II corpuscles is
greater than that of the fibres innervating the Type I clusters, the centri-
petal conduction velocity of the Type II volley is faster (by some 20-40
metres per sec.) than that of the impulses emanating from Type I corpuscles.
In summary, then, the Type II corpuscles can be regarded solely as
dynamic mechanoreceptors, whose brief, high-velocity discharges signal
joint acceleration and deceleration.
* Some previous workers have regarded these corpuscles as modified forms of the
Vater-Pacinian corpuscle-but, for reasons discussed elsewhere (Wyke, 1967), we
do:not agree with this homology (see also Fitzgerald, 1962).
33
 BARRY WYKE

Tjpe III receptors

Whereas the Type t and Type 11 corpuscles are joint capsule receptors,
the Type III corpuscles-an example of which is shown in Figure 4c-are
confined to the joint ligaments. They are the largest of the articular
corpuscles, and are identical structurally with the tendon organs of Golgi-
of which they appear to be the articular homologue. As can be seen in
Figure 4c, each Type III corpuscle is a fusiform endorgan applied longi-
tudinally to the external surfaces of the joint ligaments, usually near their
bony attachments; and it consists of a filmy connective tissue capsule en-
closing a mass of densely arborizing nerve filaments derived from a large,
Group 1 myelinated parent axon that may be up to 17p in diameter.
A few of these corpuscles are located on all the extrinsic (that is, the
collateral) ligaments of the limb and spinal apophyseal joints, and on
intrinsic joint ligaments such as the cruciate ligaments in the knee joint.
Some are also present in relation to the lateral ligament of the temporo-
mandibular joint; but they are absent from the longitudinal ligaments of
the vertebral column. The Type III corpuscles are also entirely absent
from the ossicular joints of the middle ear, and from all the laryngeal
joints.
The stimulus-response characteristics of these endorgans have not yet
been studied as thoroughly as those of the Type I and Type II corpuscles;
but the data available thus far suggest that the Type III endorgans behave
as high-threshold, slowly adapting mechanoreceptors-as do the Golgi
endorgans in the tendons related to joints. Thus they are completely
inactive in immobile joints, and only become active towards the extremes
of joint movement-that is, when considerable stresses are generated in the
joint ligaments. In these circumstances, the Type III corpuscles emit
a stream of impulses that travel centripetally at high velocity in the large
diameter afferent fibres in the articular nerves; and the frequency of this
discharge adapts only very slowly (over many seconds), if the extreme
joint displacement be maintained.
Type IV receptors
The Type IV category of articular receptors embraces the non-corpus-
cular nerve endings in the joint tissues, being represented by lattice-like
plexuses (Fig. 4d) and free nerve endings (Fig. 4e). These terminations
are derived from the smallest (Group III) afferent fibres in the articular
nerves-some of which (those between 2g and 5p in diameter) are thinly
myelinated, whilst the remainder (less than 2g in diameter) are unmye-
linated.
The plexus or network system of terminals, an example ofwhich is shown
in Figure 4d, is prominent in the limb, spinal apophyseal and temporo-
mandibular joints, in each of which it is distributed throughout the fibrous
capsule and adjacent periosteum, the articular fat pads (both external and
internal), and the adventitial sheaths of the articular blood vessels. In the
34
 THE NEUROLOGY OF JOINTS
tissues of these joints, free nerve endings are sparse, being confined largely
to the intrinsic and extrinsic joint ligaments, as shown in Figure 4e. In the
capsules of the laryngeal and ossicular joints, however, the network
system is less well developed, and there are then many free nerve endings
distributed throughout the fibrous joint capsule.
This Type IV category of ending constitutes the pain receptor system of
the articular tissues. As such, the plexuses and free nerve endings are
entirely inactive in normal circumstances; but they become active when
the articular tissues containing this type of ending are subjected to marked
mechanical deformation, or to direct mechanical or chemical irritation.
In this connection, it should be emphasized that this variety of receptor
ending is entirely absent from the synovial lining of every joint that has
been examined, and is also lacking from the menisci present in the knee
and temporo-mandibular joints, and from the intervertebral discs. There
is no mechanism, then, whereby articular pain can arise directly from the
synovial tissue or menisci in any joint; and surgical removal of synovial
tissue or joint menisci likewise does not involve removal of pain-sensitive
articular tissues.
FUNCTIONS OF ARTICULAR RECEPTOR SYSTEMS
Having described the general pattern of innervation of the joints of the
body-which appears to be essentially similar in all mammalian joint
systems-I should like to devote the remainder of this lecture to a con-
sideration of the functional significance of this articular innervation.
Articular kinaesthetic sensation
Although Sherrington and his colleagues customarily included the
joints within the system of proprioceptive inputs that influence muscle
tone, his interests in this respect were largely in the receptors in the muscles
themselves. For this reason, it was for long believed that the muscular-
and to a lesser extent the cutaneous-mechanoreceptors were the principal
reflex regulators of striated muscle tone at a segmental level, and the major
contributors to postural sensation.
In recent years, however, it has become apparent that mechanoreceptors
in the articular tissues make substantial contributions to the perception
of joint posture and movement-a contribution that is not made by the
muscle receptors. This suggestion-that receptors in the joints might be
significant contributors to kinaesthetic sensation-was originally made
at the end of the last century by Goldscheider (1898); but it is the con-
siderable amount of recent work in experimental psychology and neurology
over the last decade that has provided ample confirmation of their im-
portance in this respect-so much so, that the articular mechanoreceptors
can now be regarded as the major contributors to postural and kinaesthe-
tic sensation, supplementary to the information provided by vision and
from the skin.
35
 BARRY WYKE
Articular mechanoreceptor (arthrokinetic) reflexes
Less adequately delineated, however, is the possible r6le of the articular
receptors in the reflex regulation of muscle tone; and so it is to this aspect
of their function that we have particularly directed our attention. In
the remainder of this lecture, then, I should like to outline the results of
some of the experimental and clinical studies that we have made in order
to throw light on the reflex significance of the articular receptor systems
that I have just described.
Previous experiments have indicated that the discharges from the Type It
endorgans-which, you will recall, behave as low-threshold, rapidly
adapting mechanoreceptors-are transient, occurring only at the onset or
cessation of movement. Our investigations have confirmed that this is
indeed so; but they have further revealed that these transient discharges
from the Type II receptors are responsible for provoking brief reflex
changes in the tone of the muscles operating over the joints being moved-
changes that may be facilitatory or inhibitory in nature, and which are
reciprocally co-ordinated between individual muscle groups.
One of the sites in which this effect can be conveniently demonstrated is
the larynx, where the joints (with the sole exception of the crico-arytenoid
joint) contain only Type II mechanoreceptors. If we take a joint such as
the cricothyroid joint, which we know to be provided solely with Type II
receptors, it can be shown (Figs. 5, 6) that passive movements of this joint
give rise to very short-lived, reflexly co-ordinated changes in the tone of
the intrinsic muscles of the larynx.
Figure 5a shows portion of an electromyogram recorded from the
detached left cricothyroid muscle, during the performance of passive
movements of the isolated left cricothyroid joint. It is apparent from this
tracing that the onset and cessation of joint displacement is, in each in-
stance, associated with a very brief burst of motor unit potentials in the
muscle, the time course of which is compatible with their being provoked
by the evanescent discharge of the Type II mechanoreceptors in the moving
joint capsule. Furthermore, as Figure Sb shows, movements of the
cricothyroid joint in appropriate directions may produce equally short-
lived changes in the tone of the other intrinsic muscles of the larynx that
are reciprocally co-ordinated. Thus, as can be seen in Figure Sb, antero-
medial displacement of the cricothyroid joint produces a brief increase in
motor unit activity in the thyro-arytenoid muscle (upper tracing), that is
accompanied by an equally brief inhibition of motor unit activity in its
antagonist posterior crico-arytenoid muscle (lower tracing).
That these changes in motor unit activity do in fact arise reflexly in
response to the discharge of mechanoreceptors located in the moving joint
capsule can be demonstrated in four types of control experiment.
First, we have shown that identical reflex responses can be provoked in
these same muscles by direct electrical stimulation (with appropriate
36
 THE NEUROLOGY OF JOINTS
parameters) of the articular nerves supplying the cricothyroid joint.
Secondly, we have shown that the reflex muscular responses to cricothyroid
joint movement (such as are depicted in Figure 5) can be totally and per-
manently abolished by section of these same nerves. Thirdly, if the
receptor nerve endings in the joint capsule be destroyed by direct electro-
coagulation, then all the reflex changes in laryngeal muscle tone that
otherwise occur when the joint is moved are abolished and this abolition
is permanent. On the other hand, reversible abolition of the rapidly

'
(a) Reproduced by couirtesy of Ann. Otol. Rhinol. Laryngol.
Fig. 5. Electromyograms of articular reflexes in the intrinsic muscles of the
cat's larynx, recorded from pairs of needle electrodes inserted directly into the
muscles. (a) Rapidly adapting motor unit discharges provoked in the detached
left cricothyroid muscle at the onset and cessation of passive caudal displacement
of the isolated ipsilateral cricothyroid joint. The interrupted line indicates a
period of 60 sec. during which constant joint displacement was maintained. (From
Kirchner and Wyke, 1965d.) (b) Reciprocally co-ordinated, rapidly adapting
articular reflex responses in the thyro-arytenoid muscle (above) and posterior
crico-arytenoid muscle (below), displayed in simultaneous recordings during
antero-medial displacement of the isolated ipsilateral crico-thyroid joint. Note
that, immediately following the joint movement, brief facilitation of motor unit
activity in the (adductor) thyro-arytenoid muscle is accompanied by transient in-
hibition of activity in the (abductor) posterior crico-arytenoid muscle. Note also
that slowly adapting changes in motor unit activity are absent in each of the
tracings in (a) and (b).
adapting reflex responses in the laryngeal muscles can be produced by
infiltration of the capsule of the cricothyroid joint with a very small
amount of a 1 per cent solution of Lignocaine hydrochloride-a potent
local anaesthetic agent that has a short-lived action. Thus, in Figure 6
you can see that the rapidly adapting motor unit response that is provoked
reflexly in the thyro-arytenoid muscle by movement of the cricothyroid
joint (upper tracing) is abolished within six minutes of infiltrating the joint
capsule with this local anaesthetic agent (second tracing). Fifty minutes
37
 BARRY WYKE

later, however, as the effects of the local anaesthetic agent begin to wear off,
the motor unit response to joint movement can be seen to be re-emerging
(third tracing); and this process continues, so that after the lapse of about
one hour the motor unit potentials are well on the way back to their
original pattern (fourth tracing).

Reproduced by courtesy of Ann. Otol. Rhinol. Laryngol.

Fig. 6. Reversible suppression of articular reflex responses in the thyro-arytenoid
muscle to passive movement of the isolated ipsilateral cricothyroid joint by local
anaesthesia of the joint capsule with 0.1 ml. of 1 per cent Lignocaine hydrochloride
solution. Note that the rapidly adapting motor unit response to caudal joint
displacement seen in the upper electromyogram is absent in the second tracing: the
third and fourth tracings show the gradual return of the motor unit responses as the
effect of the anaesthetic agent on the nerve endings in the joint capsule wanes.
(From Kirchner and Wyke, 1965d.)
In our opinion, this type of experimental sequence-which we have
also applied to the limb and temporo-mandibular joints (vide infra)-lends
powerful support to the view that the Type 11 receptors are in fact rapidly
adapting mechanoreceptors that are provoked into transient discharges
at the onset and cessation of joint movement. Further, such experiments
reveal that these brief discharges reflexly provoke short-lived bursts of
38
 THE NEUROLOGY OF JOINTS
motor unit activity in some of the muscles related to the joint, with coin-
cident short-lived inhibition in other muscles functionally related to the
same joint.
Thus the Type II receptors may be regarded as behaving (in most sites,
at least) as " booster" mechanoreceptors, evoking supplementary motor
unit activity transiently in the prime movers of a joint at the onset of
movement (with coincident brief inhibition of their antagonists) in order to
overcome the inertia of the immobile parts; but once the joint is in motion
their activity ceases, until the parts are brought to rest once more. In the
light of this information it will be apparent that the Type IL articular
mechanoreceptors can make no contribution to the perception of static
joint position; nor is it likely, in view of the very brief duration of their
afferent discharge, that they make any significant contribution to per-
ception of joint movement. In short, the Type II mechanoreceptors
appear to be purely reflexogenic.
In the joints of the body other than the larynx and the middle ear, how-
ever, the reflex operations of the rapidly adapting Type 1I mechanorecep-
tors are supplemented, in normal circumstances, by those of the slowly
adapting Type I mechanoreceptors; and I should like to illustrate these
combined effects in two different joint systems-beginning with the tem-
poro-mandibular joints.
As Figure 7 shows, stimulation of the Type It receptors in one temporo-
mandibular joint by movement of the ipsilateral half of the mandible
provokes transient reflex changes in motor unit activity in the related
masticatory muscles at the onset of jaw movement, similar to those I have
demonstrated in the laryngeal muscles. For example, in the upper part
of Figure 7 you can see that slow centric opening of the mouth provokes,
in the isolated ipsilateral temporalis muscle, a transient discharge of motor
units at the beginning of the jaw movement. The bottom half of Figure 7
shows, however, that centric closure of the mouth provokes a slowly
adapting (and fluctuating) increase in motor unit activity in the same
muscle, that persists throughout the time that the jaw is kept closed but
stops when the mouth is re-opened. In this case, then, passive movements
of the temporo-mandibular joint in appropriate directions give rise both
to rapidly adapting and to slowly adapting reflex changes in the tone of
the masticatory musculature-and this correlates with our histological
observations that the temporo-mandibular joint capsule contains a mixture
of Type II and Type I mechanoreceptors. That these reflex effects arise
from mechanoreceptors located in the capsule of the temporo-mandibular
joint has been confirmed by an experimental procedural sequence (in-
volving articular nerve stimulation and section, and electrocoagulation and
local anaesthesia of the joint capsule) similar to that already described in
respect of the laryngeal joints.
This group of experiments suggests, then, that in those joints equipped
both with Type I and Type II mechanoreceptors, the Type I receptors
39
joints.

~ ~ ~ centri19v BARRY WYKE

exercise a continuous (or tonic) influence over the reflex regulation of the
tone of the muscles operating over the joint in question-in contrast (and
supplementary) to the discontinuous (or phasic) influence exerted by the
Type II receptors. This combination of influences is particularly im-
portant in respect of the mechanoreceptors located in the capsules of the
limb joints; and one further set of experiments will serve to demonstrate
the co-ordinated reflex effects of the mixture of rapidly adapting Type 11
and slowly adapting Type I mechanoreceptors that is present in all the limb

These effects can be illustrated in respect of any limb joint; but the
one I have chosen for present purposes is the ankle joint. For example
(Fig. 8a), an electromyogram of the tenotomized tibialis anterior muscle
during passive plantarflexion of the related foot shows that, as the move-

Open
TZM~ALE.

~'lsdOpeuil1-
Centric~~~~~~~~Cnti

I sec.

Fig. 7. Electromyograms of temporo-mandibular articular reflexes in the

detached temporalis muscle of the cat, provoked by passive movements of the iso-
lated ipsilateral ramus of the mandible. Both tracings are from the same prepara-
tion, with bipolar needle electrodes inserted directly into the muscle: all move-
ments were made without occlusal contact of the teeth. The upper tracing, and
the right-hand end of the lower tracing, shows rapidly adapting motor unit responses
provoked by passive opening of the mouth: the principal part of the lower tracing
shows the fluctuating, slowly adapting responses to passive closure of the mouth.

ment begins, there is a brief, high-amplitude discharge of motor units

that dies away within about two seconds, melting into a more prolonged
discharge of motor units that persists-but with a slow diminution in
frequency-as long as the foot is held in the plantarflexed position. Con-
versely, as Figure 8b shows, passive dorsiflexion of the same foot provokes
a similar dual pattern of discharge in the tenotomized gastrocnemius
muscle. As our histological investigations show that the capsule of the
ankle joint contains both Type I and Type II receptors, this dual pattern
of muscle reflex response to ankle joint movement is to be expected; and
again, in the light of the data already presented, it seems reasonable to
attribute the initial rapidly adapting motor unit discharge to the reflex
effects of Type II receptor activity; whilst the slowly adapting, lower-
frequency discharge that persists during maintenance of the joint in its
40
 THE NEUROLOGY OF JOINTS
new position can be attributed to the reflex effects of the Type I receptor
discharge.
Proof that these reflexes are in fact derived from mechanoreceptors in
the capsule of the ankle joint is provided by the same experimental pro-
cedures as before, of which Figure 9 provides a typical illustration. Thus,
as Figure 9 shows, electrocoagulation of the ankle joint capsule per-
manently abolishes the rapidly and slowly adapting motor unit responses
in the tenotomized tibialis anterior muscle during plantarflexion of the
foot. Again, as in the experiments on the larynx (Fig. 6) and with the
temporo-mandibular joints, the motor unit responses in the leg muscles

(a) , 00jIOOpV
2 Sec.

(b) , O#
2'Sec.
Fig. 8. Electromyograms of articular reflex responses in the leg muscles of the
cat to passive movements of the ipsilateral ankle joint. Prior to the recordings, a
cuff of skin was removed from around the joint region, and all tendons operating
over the joint were divided and freed. (a) Rapidly and slowly adapting motor unit
responses in the tenotomized tibialis anterior muscle to plantarflexion of the foot.
(b) Rapidly and slowly adapting motor unit responses in the tenotomized gastroc-
nemius muscle to dorsiflexion of the foot.
to passive movement of the ankle are temporarily abolished by infiltration
of the capsule of the joint with a small quantity of local anaesthetic
solution, and permanently abolished by articular neurectomy.
Finally, it should perhaps be pointed out that direct mechanical stimu-
lation of the capsule of any of the joints is capable of provoking similar
reflex responses in the related muscles. Thus, as Figure 10 shows, direct
gentle compression of the capsule of the ankle joint provokes both rapidly
and slowly adapting increases in the motor unit activity of the related
gastrocnemius muscle (a)-and these responses can no longer be obtained
after electrocoagulation of the nerve endings in the joint capsule (b).
As a result of these experimental studies, I hope you will allow that it is
reasonable to propose that the mechanoreceptors in the capsules of all the
41
 BARRY WYKE
joints make significant contributions to the reflex regulation of the tone
of the muscles related to those joints-provoking either facilitation or
inhibition during joint displacement, depending upon the direction and
amplitude of the movement of the joint in question. At the onset of
movement, the Type II receptors in the joint capsule and fat pads are
provoked into a brief episode of high-velocity impulse discharge that gives
rise reflexly to a short-lived, reciprocally co-ordinated alteration in motor
Before Electrocoogulation Of Toint Capsule

(a) L L.JlOO)1V
2 Sec.

After Electrocoagulation Of 3oinl Capsule

(b) . , 10OO)JV
2 Sec.
Fig. 9. Abolition of articular reflex responses in the leg muscles to passive move-
ment of the ankle joint by electrocoagulation of the joint capsule. Both electro-
myograms were recorded from the same muscle (tibialis anterior). (a) Motor unit
responses evoked in the tenotomized tibialis anterior muscle by passive plantar-
flexion of the ipsilateral foot. (b) Following capsular electrocoagulation, the motor
unit responses are absent. The myotatic reflex (not illustrated) was still elicitable
in the muscle.
unit activity in the related muscles. At the same time, and throughout the
whole course of joint movement, the Type I receptors continuously con-
tribute a fluctuating discharge whose precise frequency varies with the
degree and velocity of joint displacement; and this gives rise reflexly to
an equally continuous but fluctuating alteration-involving either facilita-
tion or inhibition-in the activity of the motor units in the muscles oper-
ating over the joint. The Type III receptors, which (as I have mentioned)
are present only in the joint ligaments, are not provoked into activity in
the circumstances of experiments such as those I have described-unless
the joint manipulations produce extreme degrees of joint displacement:
42
 THE NEUROLOGY OF JOINTS
then, and only then, do these high threshold receptors discharge, to pro-
duce reflex inhibition of motor unit activity in the muscles operating over
the joint. The Type IV pain receptors in the articular tissues are likewise
not activated in these experiments; but other experiments that we have
performed show that they can be-by sufficiently intense irritation of the
joint capsule, fat pads or ligaments-and their discharge then provokes
intense non-adapting motor unit responses simultaneously in all the
muscles related to the joint, as well as in more remote muscles elsewhere
in the body.
Compression Of Joint Capsule Before Electrocoagulation

(a) on Off,,f
ff

2 Sec. I

Compression Of Joint Capsule After Electrocoagulotion

on off
(b) . 10gO,,#
2 Sec.
Fig. 10. Evocation of articular reflex responses in the leg muscles by direct
mechanical stress applied to the capsule of the ipsilateral ankle joint. Both
electromyograms were recorded from the same tenotomized muscle (gastrocnemius).
(a) Rapidly and slowly adapting motor unit discharges provoked by direct gentle
compression of the capsule of the ipsilateral ankle joint (maximal posteriorly).
Note that the pattern of discharge is similar to that evoked in the gastrocnemius
muscle when the posterior capsule of the ankle joint is stretched by passive dorsi-
flexion of the foot (see Fig. 8b). (b) Following electrocoagulation of the joint
capsule, similar compression no longer evokes any motor unit responses. (The
increased amplitude of the activity throughout record (b) is due to the fact that
general anaesthesia was allowed to decrease between (a) and (b), so that even slight
reflex effects could be detected, if present.)

The precise nature of the reflex pathway through which these effects are
accomplished is not yet determined; but a variety of incidental observa-
tions that we have made in the course of our experiments-observations
that I cannot detail here-suggests to us that the articular mechanoreceptor
(i.e. the arthrokinetic) reflex pathway is polysynaptic, and that it projects
to gamma (fusimotor) rather than to alpha motoneurones. If this proposal
can be substantiated in our later experiments, then the articular mechano-
receptors can be added to the cutaneous mechanoreceptors in the segmental
reflex systems driving the gamma motoneurone loops.
43
 BARRY WYKE
In the light of the observations that we have made to date, however-
and of the information now available from other laboratories-it seems
likely, then, that the Type I mechanoreceptors contribute reflexly to the
maintenance of tone in the muscles of the limbs, spine and jaws at rest,
and to the co-ordinated reflex regulation of isotonic and isometric changes
in the tone of those muscles when movements are made, or attempted;
and in addition, these same receptors contribute to conscious perception
of static joint position and of dynamic joint movement-that is to say, to
kinaesthesis. In contrast, the Type II and Type III mechanoreceptors are
reflexogenic only, the former provoking transient " booster" changes in
the tone of the muscles operating over a joint during the actual period of
acceleration or deceleration of the articulated bones, whilst the latter act
as " brakes " to limit excessive joint displacement.
THE CLINICAL SIGNIFICANCE OF ARTHROKINETIC REFLEXES
It may rightly be argued, however, that whilst these observations may be
pertinent to the experimental situations on which they are based, they do
not provide definitive evidence that arthrokinetic reflexes are of practical
significance in the circumstances of everyday life. In order to investigate
their possible significance in this respect, then, we have undertaken a
further set of experimental and clinical investigations, of which I should
like to show two examples as a conclusion to this lecture.
If the functional proposals I have made thus far are to be regarded as
applicable to the normal living subject, it should follow that interruption
of the flow of impulses from the mechanoreceptors in a joint capsule into
the central nervous system should result in clinically evident disturbances
of perception of joint position and movement, and of re1texes of posture
and gait. That this is indeed so can be shown in two ways.
First, the centripetal flow of such mechanoreceptor afferent impulses can
be interrupted by surgical denervation of a particular joint in animals, and
the subsequent behaviour of the animals can be studied for prolonged
periods, in recovery experiments. Secondly, in animals and in man, it is
possible to demonstrate, by appropriate methods of testing, that destruc-
tion of receptor endings in the capsule of a joint gives rise to persisting
abnormalities of kinaesthesis and of postural reflexes.
The effects of articular neurectomy
The first of these situations has been examined by carrying out aseptic
partial neurectomy of one knee joint in a series of cats, followed by studies
of the postural reflexes and behaviour of the experimental animals there-
after, for periods of up to two years.
In some of these animals, the posterior articular nerve to the left knee
joint was divided in the popliteal fossa; and in a second series of animals,
the medial articular nerve to the left knee joint was divided in the front o,
the thigh.
44
 THE NEUROLOGY OF JOINTS
The resulting abnormalities of postural reflex behaviour that persist
after these two operations, and which are not demonstrable in control
animals subjected to comparable dummy operations, are summarized in
Table III. From this Table, in which plus symbols indicate the severity of
the post-operative disturbances, you can see that both posterior and medial
articular neurectomy result in significant and persistent changes in certain
aspects of postural reflex behaviour. I do not propose to go into the
detailed analysis of these findings here (see Freeman and Wyke, 1966); in-
stead, I should simply like to show you two examples of the abnormal
reflexes that develop following this type of articular neurectomy.
TABLE III
REFLEX EFFECTS OF ARTICULAR NEURECTOMY
Behaviour Posterior Medial
neurectomy neurectomy
Stance .. + 0
Reaching .. o o
Gait .. .. + +
Jumping .. + +
Bar walking.. + + +
Posterior neurectomy Medial neurectomy
Reflex Ipsi- Contra- Ipsi- Contra-
lateral lateral lateral lateral
Preening .. .. o o o o
Knee-jerks .. .. o o o o
Withdrawal .. o o o o
Suspension .. ++ o + o
Stutz.. .. .. +++ o +
Placing .. ..+++ o +++ o
Crossed extensor . . o - ++
Figure 11 depicts two cats, in each of which the posterior articular nerve
supplying the left knee joint has been divided. You will notice that the
animal in the upper illustration keeps its left hind leg in an abnormal
posture while eating-the limb being externally rotated, and the knee
flexed, to an extent greater than in the opposite normal leg. Likewise, in
the animal in the lower illustration, you can see that the posture of the
left leg when the animal is walking is abnormal-the whole of the foot
being placed on the walking surface, instead of just the toe pads. Further-
more, as the data in Table III indicate, these animals are generally un-
steady in their gait, and in jumping; and furthermore, they clearly behave
as if they had a persisting kinaesthetic sensory deficit related to the left
hind leg-in that when walking, for instance, they frequently stop and
shake the limb, and turn round to look at it.*
The effects of articular receptor damage
In our view, this series of experiments provides clear evidence that dis-
* In relation to the above observations, it should be noted that none of these animals
developed pathological changes in the structure of the articular surfaces, or of the
connective tissues of the joint capsule-although these were sought when the animals
were destroyed at various intervals after operation.
45
 BARRY WYKE
tortion of the normal centripetal flow of mechanoreceptor afferent im-
pulses from a joint produces significant impairment of reflex muscular
behaviour, as well as of kinaesthesis. Similar disturbances of postural
reflex behaviour can be detected also in human beings, when the receptor
nerve endings in a joint are damaged or diseased. As an example of this,
Figure 12 shows the results of some preliminary clinical postural studies
carried out in collaboration with Mr. Michael Freeman.

ii: ~~.
............................................
....... ia

(a)

Reproduced by courtesy of British Journal of Surgery.

Fig. 11. Persisting postural abnormalities in the cat following neurectomy of the
posterior articular nerve to the left knee joint. (a) Abnormal static posture of the
left hind leg during feeding (description in text). (b) Abnormal dynamic posture
of the left hind leg during walking (description in text). (From Freeman and
Wyke, 1966.)
The boy shown in Figure 12 had sustained an injury to his right ankle
joint (indicated by the black bar in the photographs) whilst playing foot-
ball some months previously, that resulted in damage to the lateral cap-
sular tissues of that joint. Although he made what appeared to be a
perfectly satisfactory recovery from this injury, and was completely free
of pain, he subsequently complained that his right leg felt unstable; and
for this reason, we carried out a systematic examination of his post-ural
reflexes. This was done with a special technique that we have developed
46
 (a)

(b)
*:~ ~ ~ . .:
THE NEUROLOGY OF JOINTS

V~~~~~~~~~~~~~~~~~
i :

Fig. 12. Persisting postural abnormalities in man resulting from disturbance of

articular reflexes operated from the ankle joint, following injury to the lateral
capsule of the right joint. The two illustrations on the left show the normal posture
when standing (with the eyes closed) on the unaffected left foot, with the arms at
the sides (a) and elevated (b). The two illustrations on the right show the differ-
ences when standing on the affected right foot. Partial description in text: in
addition, note (i) Depression of shoulders when standing on right foot, indicated
by a decrease of 100 in the obtuse angle between the shoulder tangents drawn in on
the left and right of Figure 12a. (ii) Adduction and supination of ipsilateral arm,
and extension of ipsilateral fingers, with flexion of contralateral elbow and fingers,
when standing on right foot (compare Bl and B2). (iii) External rotation and
abduction of contralateral leg, when standing on right foot (compare Cl and C2).
(iv) External rotation of ipsilateral leg, and flexion of ipsilateral toes, when standing
on right foot (compare Dl and D2). (v) Partial extension of contralateral knee, and
abduction of contralateral thigh, when standing on right foot (compare El and E2).
(vi) Abduction of ipsilateral outstretched arm, and adduction and elevation of
contralateral arm, when standing on right foot (compare Xl and Y2; Yl and X2).

47
 BARRY WYKE
for this purpose in our laboratory, in which postural electromyography is
combined with electronic flash photography of the patient during assump-
tion of a variety of standardized postures.
As you can see on the left of Figures 12a and 12b, when the boy stands on
his normal leg with his eyes closed, and his arms at his sides (Fig. 12a) or
elevated (Fig. 12b), the vertical axis of his body (represented by the dotted
line A1) is very close to the true vertical axis. The other interrupted line
represents the axis of his supporting leg; and I would like to draw your
attention particularly to the angle (of 6°) that this supporting axis normally
makes with the body axis. On the other hand, when the boy stands on
the right foot-the side that had been injured-you will notice (on the
right of Figures 12a and 12b) that the body axis (A2) tilts away from the
true vertical axis (by some 70 in fact) and moves towards the axis of the
supporting leg, so that the normal 6° angle between the body axis and the
axis of the supporting leg is reduced (for instance, by 50 per cent in
Fig. 12a). At the same time, the axis of the supporting leg deviates away
from its normal angle with the true vertical axis, by about 3°.
In addition to these particular disturbances, there are a number of other
postural reflex changes to be seen in this boy when he is standing on his
affected leg, some of which are indicated by the numbered arrows in the
illustrations and noted in the legend to the figure; but I cannot analyse
them further here. I simply wish to show this case, in order to lend weight
to the hypothesis advanced earlier-namely, that when injury to or disease
of a joint is such as to cause degeneration of the nerve endings in its
tissues, then this may result in postural reflex abnormalities (as well as
kinaesthetic disturbances) that are of clinical significance.
CONCLUSION
Taken in conjunction with the animal experiments that I have described
to you, and in the light of work that has been done in other laboratories to
date, these observations can leave little doubt that the mechanoreceptors
located in the fibrous capsules, fat pads and ligaments of joints are of con-
siderable importance in the circumstances of everyday life-not only in
respect of their contribution to perceptual awareness of static joint position
and of joint movement, but also in the reciprocal reflex regulation of
muscle tone in posture and movement. In the face of this evidence, I
would suggest that serious attention must now be given by clinicians-
particularly by orthopaedic and neurological surgeons, and by practitioners
of physical medicine-to the role of articular reflexes in the regulation of
normal and abnormal muscle tone, in addition to the role of the articular
receptors in kinaesthetic perception. I hope you will regard it as reason-
able to assert that articular reflexes must now assume a position of greater
importance in the physiology and pathology of movement at every joint in
the body than has been the case in the past; and that in many branches of
surgery-ranging from otology and laryngology, through orthodontics, to
48
 THE NEUROLOGY OF JOINTS

orthopaedic and neurological surgery-the arthrokinetic reflexes must be

considered as matters of moment in clinical practice. Certainly, a great
deal still remains to be done in this field, especially in defining more pre-
cisely the contribution of disordered articular reflexes to particular clinical
abnormalities of posture and movement in the various joint systems of the
body; but it is my firm belief that sufficient evidence is now available at
least to suggest that such an approach should be made. We hope, by our
continued efforts in the Neurological Laboratory of this College, to make
some small contribution to the further development of the subject of
articular neurology-and I should like to thank the President and Council
of the College for the opportunity of presenting this epitome of our pro-
gress to date.
ACKNOWLEDGEMENTS
May I conclude by thanking the Council of the College for the privilege
of delivering this Arris and Gale Lecture. The honour thus conferred
should not, however, be regarded as personal, but rather as a corporate
tribute to the work of the Research Fellows whose names are listed in
Table IV, the fruits of whose labours in the Neurological Laboratory in
TABLE IV
RESEARCH FELLOWS IN THE NEUROLOGICAL LABORATORY
Department of Applied Physiology, Royal College of Surgeons of England
Michael A. R. Freeman, M.A., M.D., F.R.C.S.
(Department of Orthopaedic Surgery, Westminster Hospital, London).
John A. Kirchner, M.D., M.S., F.A.C.S.
(Department of Otolaryngology, Yale University School of Medicine).
Bernard Greenfield, B.D.S., F.D.S.R.C.S.
(University College Hospital School of Dentistry, London).
Brian Reeves, M.B., B.S., F.R.C.S.
(Leverhulme Research Fellow in Orthopaedics).
M. A. Abo-El-Enein, M.B., Ch.B., Dip. Anat.
(Department of Anatomy, Al-Azhar University, Cairo).
the Department of Applied Physiology I have attempted to summarize
in this Lecture. I should also like to take this opportunity of acknowledg-
ing publicly the generosity of the College in constructing the specially
designed laboratory suite in which our researches are conducted; of the
Postgraduate Medical Federation of the University of London for pro-
viding a capital grant for our equipment; and of the Camilla Samuel
Fund for a recent grant to provide supplementary laboratory staff. Last,
but by no means least, I should like to pay tribute to the facilities afforded
us by Professor David Slome, and to the devoted technical skill of Mr.
S. P. Steward* and Mr. Johanna Gitau, without which the work I have
described would have been impossible.
* Mr. Steward recently retired from the staff in November 1965, with the remarkable
record of 53 years of continuous service in the College.
49
 BARRY WYKE
REFERENCES
Most of these publications are from the Neurological Laboratory, Royal College of
Surgeons of England.
ABO-EL-ENEIN, M. A., and WYKE, B. D. (1966) Nature, 209, 683.
BARNETT, C. H., DAVIES, D. V., and MACCONAILL, M. A. (1961) Synovial Joints. Their
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(1965) Nature, 207, 196.
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(1964c) Nature, 202, 600.
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(1965c) Trans. Ann. Meet. Amer. Laryngol. Assoc.

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(1965d) Ann. Otol. Rhinol. Laryngol. 74, 749.
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(1962) Ann. Roy. Coll. Surg. Engl. 30, 135.
(1965) In General Anaesthesia. 2nd ed. Vol. 1, p. 157. Ed. by F. T.
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(1967a) J. Roy. Coll. Surg. Irel. (in the press).
(1967b) Principles of General Neurology. Amsterdam, Elsevier (in the
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(1967c) Brit. J. Dis. Communic. 2, 1.
(1967 d) Studies in the Neurology ofJoints. Vol. 1. The Linib and Spinal
Joints. Amsterdam, Elsevier (to be published).
and GREENFIELD, B. E. (1967) Studies in the Neurology of Joints. Vol. III.
The Temporo-mandibular Joints. Amsterdam, Elsevier (to be published).
and KIRCHNER, J. A. (1967) Studies in the Neurology of Joints. Vol. II.
The Laryngeal and Ossicular Joints. Amsterdam, Elsevier (to be published).

A comprehensive bibliography of the subject of articular neurology is provided in the

references appended to these publications.
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