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Innervation of the eye.

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Innervation of the Eye’
J. C. BEATIE AND DONALD L. STILWELL, JR.*
Department o f Anatomy, Stanford School of Medicine, Stanford University,
Stanford, California
With the exception of the retina all
nervous tissue in the eye orginates in the
ciliary nerves. A review of the literature
reveals a lack of accurate and complete
descriptions of the sources and distribution
of these nerves. Agababow (’12) described
their distribution but included some factual
errors. There has been no comparable
work in recent times incorporating investigations made since then.
Zander and Weddell (’51), using methylene blue, demonstrated the innervation
of the cornea and reviewed the literature
to that time. Earlier Cohnheim (1867),
Hoyer (1877), Dogie1 (1890,1891), Attias
(’12), Ernyei (’34), and Boeke (’35) used
various techniques to compare the corneal
nerves in several species. In 1910 Virchow
reviewed the literature to his time and
consolidated the work of previous authors.
Zander and Weddell’s work surpasses all
others in completeness and accuracy, and
ought to replace Attias’ as the standard
reference work on the subject.
Langworthy and Ortega (’43) reviewed
the literature concerning iridial nerves,
and using methylene blue demonstrated
the essential features of their distribution.
On physiological and theoretical grounds
they deny the existence of a dilator pupillae
muscle, but describe a portion of the
sphincter muscle which passes radially
from the pupillary margin into the substance of the ciliary body. They saw fine
nerve fibers terminating in this portion of
the “sphincter muscle,” similar to the
nerves in the circularly directed muscle
fibers. Arnold (1863) was among the first
to describe the iridial innervation, and
provided an accurate drawing of the distribution of larger nerve bundles. Pause
(1877), Meyer (1879), Hosch (1891) and
Kirpitschowa-Leontowitsch(‘1 l), used various techniques in several species without
adding significantly to an understanding
of these nerves. Boeke (’33), using silver
techniques, studied the iridial nerves with
results which suffer from that technique’s
failures.
Genis-Galvez (’57) related the fine features of the ciliary muscle innervation;
Boeke (’33) and Hirano (’41) used silver
techniques to demonstrate a “vegetative
syncitium” in this muscle.
Scleral nerves were described by Bach
(1892), and Smirnow (’00). Such Stmctures as end plates, trophic endings and a
terminal nerve net are described, and
said to lie in all layers of the sclera.
Choroidal nerves were described by
Schweigger (1890), Bietti (1897), and
Ernyei (’34).
In order to provide a clear, concise, and
complete description of the innervation of
the eye the study of which this paper is
the report was undertaken.
MATERIALS AND METHODS
Thirty albino rabbits two weeks to three
months old were prepared by intracarotid
injection of 300-500 ml of methylene blue
solution, ranging in concentration from
1/2% to 1/300% ; the most effective concentration was about 1/30%. The eyes
were then removed, cut in either the equatorial or sagittal plane, fixed overnight in
8% ammonium molybdate, and washed in
tap water. The retinae were removed in a
stream of water; the irises were freed at
their periphery with sharp scissors and
carried through the process as separate
specimens. Anterior and posterior halves
formed by equatorial incisions were flattened by making 4 radial incisions from
the periphery. The nasal and temporal
‘Supported in part by grants of the National
Science Foundation and San Mateo County Heart
Association.
BThe technical assistance of Max L. Millsap
and Robert 0. Johnson is gratefully acknowledged,
45
46
J. C . BEATIE AND DONALD L. STILWELL, JR.
halves from the sagittally divided eyes
were further cut to form 4 quadrants.
These specimens were then flattened between slides, dehydrated and cleared in
alcohols, xylene, and benzyl benzoate, then
stored in the dark. Some specimens are
still adequate for observation three years
after preparation.
Two eyes were stained by immersion in
0.2% osmic acid for one hour; dehydration and clearing were performed as above.
These were used to determine the extent
of myelination and to provide a picture of
the gross distribution of nerves. Fiftymicron sections were made of two eyes
stained with methylene blue and one eye
stained with Mallory azan for reference
purposes.
Summary of innervation
The ciliary nerves have sympathetic,
parasympathetic, and somatic sensory
components. The most complete general
descriptions are to be found in textbooks of
anatomy and ophthalmology. The best
works of this kind are Duke-Elder's Textbook of Ophthalmology, and Wolffs T h e
Anatomy of the Eye and Orbit. References
to specific points in other authors' works
are provided.
Sympathetic nerves. Fibers destined for
the eye arise in the lateral cell column of
the first 3-4 thoracic spinal cord segments
(Ray, '43), and travel in the cervical sympathetic trunk to the superior cervical
ganglion. Synapses are made here with
postganglionic neurons, the axons of which
pass in the internal carotid plexus to the
cavernous sinus. Here postganglionic fibers leave the plexus both as a slender
sympathetic root of the cilary ganglion and
as numerous filaments to the ophthalmic
division of the trigeminal nerve. From the
ciliary ganglion, through which these fibers pass without interruption, the postganglionic fibers proceed in 6 to 8 short
ciliary nerves to encounter the globe in an
area encircling the optic nerve. Two,
sometimes three, long ciliary nerves branch
from the nasociliary division of the ophthalmic nerve to carry the remaining sympathetic fibers to the eye. Sympathetic
fibers are said to innervate the dilator
pupillae muscle and vessels of the choroid,
ciliary body, iris, and pericorneal (limbic)
area.
Parasympathetic nerves. Preganglionic
neurons arising in the small cells of the
accessory nucleus of the oculomotor nerve
(Edinger-Westphal group, Crouch, '36)
pass through its inferior division via the
branch to the inferior oblique muscle.
From this branch the parasympathetic root
of the ciliary ganglion arises. In the ganglion synapses are made with postganglionic neurons, the axons of which pass
in the short ciliary nerves, eventually to be
distributed to the sphincter pupillae and
ciliary muscles.
Somatic sensory nerves. Sensory cell
bodies of these neurons lie in the Gasserian
ganglion; their peripheral processes arise
as simple terminals in various eye structures. These somatic fibers leave the eye in
both long and short ciliary nerves and
reach the ophthalmic nerve in its nasociliary branch. These contribute various
sensory modalities: pain, tactile, warmth,
coolness (Lele and Weddell, '55), proprioception in ciliary and pupillary changes
(Langworthy and Ortega, '43) and vasosensation. The last two seem speculative.
OBSERVATIONS
Ciliary nerves innervated only bulbar
structures, including the most anterior
bulbar conjunctivum, where an overlap
occurred with its extraocular innervation.
The eye was supplied (excepting retina)
only by 2-3 long and 6-8 short ciliary
nerves, which penetrated the sclera within
2 mm of the optic nerve.
Long ciliary nerves. (Postganglionic
sympathetic and somatic sensory fibers.)
The long ciliary nerves arose as branches
of the nasociliary nerve, and joined in
close association with the long posterior
ciliary arteries to reach the globe near the
optic nerve. These corresponding arteries
and nerves maintained a close association
throughout most of their intraocular course
(figs. 1-3). Together they pierced the
sclera in an oblique direction, attained the
suprachoroidal space, and were directed
toward the anterior pole. Within a few
millimeters of their entrance medium-sized
bundles of nerve fibers left the main ciliary
nerves, to parallel closely the accompanying artery. The long
INNERVATION OF EYE
medium-sized branch remained at a distance of less than 1 mm from the artery to
form a para-arterial arrangement. Small
bundles of these nerves encircled or entwined and followed the vessel in plexiform fashion (figs. 1, 2). From the encircling nerves individual fibers innervated
smooth muscle of the vessel wall. This
latter arrangement is referred to as a
periarterial plexus, and is believed to include both vasomotor and vasosensory fibers. At points where the vessel branched,
whether to choroid, sclera, ciliary body, iris
or pericorneal region, contributions from
both para- and periarterial nerves accompanied the daughter branch. In general
the innervation of vessels in the eye was
similar to those elsewhere in the body.
Short ciliary nerves.
(Sympathetic,
parasympathetic, and somatic sensory fibers.) Six to 8 short ciliary nerves passed
forward from the ciliary ganglion to pierce
the sclera in a small area around the optic
nerve, similar to the long ciliary nerves.
Entering the suprachoroidal space, these
nerves passed forward, branching 4 or 5
times, to participate in a loose network
with similar small branches of other long
and short ciliary nerves. No fibers from
the short ciliary nerves were seen to enter
the choroid. Its vasculature appeared to
be innervated only by long ciliary nerves.
Occasionally small fibers entered the sclera
to terminate freely (figs. 1, 5). It thus appeared that only intercommunicating
branches and an infrequent scleral fiber
emerged from short ciliary nerves posterior
to the ciliary body.
Distribution of the combined long and
short ciliary nerues. As the long ciliary
nerves passed anteriorly medium-sized
branches formed a loose connection with
nearby short ciliary nerves, as noted above,
and shown in figure 1. At the level of the
ciliary body this process of branching and
recombination had produced a total of 2030 medium-sized bundles of ciliary nerves,
whose fibers were so intermingled that
their origins were impossible to determine.
An inextricable mixture of functional nerve
components thus resulted.
a. Choroid. Arteries of the choroid
(tunica vascularis) of the rabbit, as shown
by our Berlin blue injections, are branches
of long posterior ciliary arteries, forming
47
an anastomotic network of vessels. Their
innervation was derived only from the long
ciliary nerves, through para- and periarterial arrangements shown in figures 14. All choroidal vessels, both venous and
arterial, received nerves from this source;
no contributions were observed to arise
from the short ciliary nerves. As the paraarterial nerves repeatedly divided to follow
the vascular branches, the number of
nerve fibers in each bundle decreased accordingly. In the arteriolar portion of their
course these nerves were formed of bundles
of 2-3 fibers which ran in plexiform fashion along the vessels, but in no way different from the vascular innervation elsewhere in the body. All endings were related
to arteries or veins; no free or discrete
terminals were found. Afferent signals
from the choroid may therefore be limited
to vasosensory functions. Contrary to the
findings of Agababow and others, there
were no ganglion cells observed in the
choroid.
b. Ciliary body, From the 20-30 combined long and short ciliary nerves which
reached the level of the ciliary body medium-sized branches entered its substance.
There they divided, the divisions passing
circumferentially until they combined with
a neighboring nerve, forming an arcade at
the periphery of the ciliary muscle. Small
recurrent bundles, probably misdirected
motor fibers, from the pericorneal plexus
(to be described later) also joined in fonning the arcade system (figs. 1, 10). Nerves
passed directly from the arcade into the
ciliary muscle, to break up into numerous
filaments lying parallel to the muscle fibers
(figs. 1, 9). No complicated endings such
as “rings” (Genis-Galvez, ’57) or “terminal
syncitium” (Hirano, ’41; Boeke, ’ 3 3 ) were
observed. Vasomotor nerves were similar
to those in the choroid. No typical iree
somatic sensory endings were observed, but
the presence of vasosensory endings cannot be excluded, and are assumed to be a
part of the vascular innervation, as elsewhere.
c . 17is. The nerves to the iris had a
similar, but more distal origin to those of
the ciliary body. These nerves were somewhat larger, and the iridial arcades a bit
more complex. Immediately anterior to
branches innervating the ciliary muscle
48
J. C . BEATIE AND DONALD L. STILWELL, JR.
the conjoint ciliary nerves delivered 20 or
more bundles, which by branching and recombining formed an arcade at the base of
the iris. From its distal limit the arcade
gave rise to tortuous, radially oriented
nerves which reached the pupillary margin,
there to give off circumferentially-directed
twigs to the poorly developed sphincter
pupillae muscle (figs. 1, 8). In their radial
course these nerves also delivered fibers to
the body of the iris, probably to smooth
muscle of vascular wall and dilator pupillae, as well as to freely-ending fibers on the
anterior iridial surface, probably sensory
(figs. 6 , 7). Langworthy and Ortega (’43)
described no dilator muscle; however,
Wolff (’48) clearly demonstrated the presence of a dilator pupillae.
A terminal “vegetative syncitium,” as reported by Boeke ( ’ 3 3 ) and others employing silver stains, was nowhere visible.
Specialized nerve endings were lacking;
again, there was no feature present to indicate a difference from the innervation of
smooth muscle elsewhere. Vessels of the
iris were supplied with periarterial fibers
derived both from the tortuous, radially
directed nerves and other nerves entering
with the vessels.
d. Cornea. Ciliary nerves distal to their
iridial branches were destined almost exclusively for the limbus and cornea. Figure 1 shows that terminal divisions of the
ciliary nerves consisted of 15-20 major
branches, which entered and ramified in
the substantia propria of the cornea, and
40-50 smaller twigs which recombined to
form a circumferential pericorneal plexus
at a more superficial episcleral level (figs.
1, 10).
Fibers of this pericorneal plexus (Zander
and Weddell : episcleral pericorneal plexus,
plexus paramarginalis of Attias, plexus annularis of Ranvier, Randplexus of Dogiel),
were mostly directed into the peripheral
one-third of the cornea, bundles passing
radially inward from the episcleral plane to
terminate in freely arborizing endings of
the epithelium (fig. 15). Directly superficial to the plexus, at the limbus, these
free endings were especially profuse in the
epithelium, much more numerous than in
the central cornea (fig. 14). A few fibers
of the plexus recurred into the ciliary mus-
cle (fig. 10) to complete its innervation
(most probably “misdirected motor axons),
and into the region of the tendinous insertion of extrinsic muscle, there to end
in branched, unspecialized free endings,
much as periosteal nerves are locally abundant at a tendon attachment (figs. 1, 11).
Many limbic branches of the plexus innervated the profuse vasculature of this
region (fig. 14).
The larger, major ciliary nerve branches,
presumed to be somatic sensory, proceeded
into the substance of the cornea to form a
plexiform aggregation in the substantia
propria, most highly developed immediately beneath Bowman’s membrane (figs.
1, 10). Axon groups penetrated the membrane to end freely in epithelium. Organized or encapsulated terminals were not
observed, as they are in conjunctivum, and
the axon tips were always discrete, never
syncitial (figs. 12, 13). The “rare ganglion cells” along branches of the ciliary
nerves were never encountered here or
elsewhere in the eye, as claimed by several
authors (Zander and Weddell, ’51, Agababow, ’12, Givner, ’39). Otherwise all details of the corneal innervation conformed
with Zander and Weddell’s more complete
account.
e. Extra-ocular muscles. As expected,
the motor and proprioceptive innervation
of somatic muscle was delivered by extraocular muscle nerves, not by ciliary nerves.
An increased density of free endings at
their tendinous insertion arose from
branches of the pericorneal plexus, as was
noted in the previous section (figs. 1, 11).
No inferences as to specialized function
can be made.
f . Sclera. Posterior to the ciliary body
the ciliary nerves occasionally gave off
small branches consisting of two axons at
the most, which terminated in simple, discrete free endings at the internal surface
of the sclera (figs. 1, 5). That these were
unrelated to the choroidal coat is established by the fact that they were undisturbed by lifting off the choroid. Such
endings bore no physical relation to organized structures, and it is assumed that they
subserve scleral sensation associated with
changes in external and internal global
pressures.
INNERVATION OF EYE
DISCUSSION
This investigation was undertaken to
provide in so far as possible an integrated
description of the sources and tenninations of nerves to structures of the globe,
excepting the retina. Most earlier investigations are confirmed; some are extended
or altered. This presentation is the unification of our own investigations and those
of others.
There is no evidence that the eye is innervated by nerves other than long and
short ciliary, except for the retina. Several sources are drawn upon for visceral
motor and somatic sensory innervation,
converging on the globe in the ciliary
nerves, all of which pierce the sclera within about 2 mm of the optic nerve. Almost
immediately these elements become combined in a loose network formed by the
anastomosis of ciliary nerves, so that all
nerve elements in the anterior two-thirds
of the globe are indistinguishably combined
into mixed nerves, composed of postganglionic visceral motor, somatic sensory,
and probably vasosensory fibers.
Ganglion cells. Givner (’39) describes
ganglion cells in the course of the ciliary
nerves prior to reaching the sclera, as well
as along the intra-scleral portion of their
course. Hirano (’41) and Boeke (’33) describe them in the ciliary body. Hosch
(1891) and Boeke found them in the iris
as well. Zander and Weddell (’51) picture
a ganglion cell in the episcleral pericorneaI
plexus, but state that they are rare. We
were unable to demonstrate structures that
we feel justified in calling ganglion cells,
in agreement with Ernyei (’34). In preparations employing heavy concentrations
of methylene blue, fat and smooth muscle
cells and their nuclei occasionally took up
the dye, giving an appearance similar to
the figures of other authors (fig. 3). However, there was no relationship between
such cells and nerve fibers other than contiguity. We therefore doubt the existence
of ganglion cells in any great number. If
they exist they must be considered ectopic
Gasserian, sympathetic, or ciliary ganglion
cell bodies.
Vegetative syncitium. Hirano (‘41) and
Boeke (’33) have demonstrated with silver
impregnation techniques what they term
the “vegetative syncitium” in the ciliary
49
body, consistent with findings of Stohr
(‘32) in other visceral structures. They believe that the fine individual fibers lying
among muscle fibers are interconnected by
a protoplasmic continuum. As figure 9
shows there is no such continuity demonstrable by the relatively specific methylene
blue technique. It is our impression that
the results of these authors and others who
utilize silver techniques are artifacts produced by the precipitation of silver salts on
cellular interfaces other than neurons.
Zander and Weddell have shown satisfactorily that what has been described as a
protoplasmic continuum in the corneal
nerves is but a demonstration of delicate
fibrillar structures in Schwann elements.
These fibrils may be demonstrated by
methylene blue if its specificity is lost by
improper techniques, i.e., overstaining.
Pertinent to this is a discussion by Miller
(’60) which hinges around terminals of
the autonomic nervous system in structures of the skin. The question there also
is whether a terminal syncitium exists anywhere. Those who argue the contrary, like
the present authors, utilize methylene blue,
while proponents of this view utilize silver
methods. As is stated by Miller the final
word must await utilization of the electron
microscope.
Trophic endings. Agababow (’12) describes fine endings which encompass cells
of the sclera without penetrating the substance of the cell. He denies the opinions
of authors, not specifically named but presumably Kuhne (1862), Lipmann (1869),
and Waldeyer and Izquierdo (1880), who
described nerves ending within the cytoplasm of corneal corpuscles. We are unable to place credence in a theory that such
structures as trophic nerves exist in sclera
and cornea. In the present study we found
no intracellular terminations of nerve fibers, nor any special embrace of a scleral
cell by nerve endings.
Choroid. The form of innervation of
the choroid has been described by Schweigger (lS90), Bietti (1897), and Agababow
(’12) as a wide-meshed plexus, derived
from branches of the ciliary nerves, which
contains numerous ganglion cells. Individual fibers from the plexus pass to vessels to
terminate on the vessel walls. Sensory
nerves have generally been denied. Ernyei
50
J. C. BEATIE AND DONALD L. STILWELL, JR.
(’34) studied the tissues of the eye, looking specifically for ganglion cells and the
relationship between myelinated and unmyelinated fibers. He states that no ganglion cells are to be found in any portion
of the eye, but that there are lightly myelinated fibers present in choroid which
could be interpreted either as sensory or
preganglionic autonomic. Since there are
no ganglion cells in the choroid, myelinated fibers are probably sensory. We
have demonstrated that the innervation of
the choroidal vessels is primary and not
through the intermediary of a plexus. A
‘plexiform arrangement is produced by the
arrangement of the vessels. We, too,
found no ganglion cells, and failed to find
discrete endings not associated with vasculature. From the profusion of nervi vasorum and the presence of lightly myelinated
nerves we are led to the conclusion that
sensory fibers are present in the choroid,
and from their location are probably vasosensory in nature. Their physiological significance cannot be adduced on anatomical
grounds alone.
Somatic senso y nerves. Simple, freely
arborizing sensory terminals in the cornea,
sclera, scleral insertion of extrinsic ocular
muscles, and epithelium of iris and ciliary
processes give rise to nerve fibers which
comprise the somatic sensory components
of the ciliary nerves. Other than vasosensory endings these appear to be the only
sensory terminals in eye tissues. Although
Zander and Weddell (’51) state that this
is the U S U mode
~
of free termination, they
also described terminations by “protoplasmically continuous nets formed by the fusion of daughter axons orginating from
the same parent fiber.” They denied continuity between axons of separate fibers.
A subsequent paper by Lele and Weddell
( ’ 5 6 ) stated that there are no terminal
nets, although shrinkage of the specimen
may sometimes produce this picture. We
confirm their opinion that organized terminal nerve nets and protoplasmic continuity between different axons apparently
do not exist.
Evidence in support of these anatomical
findings is supplied by the neurophysiological studies of Tower (’40). Various scleral stimuli evoked infrequent, low amplitude activity in ciliary nerves, indicating
scanty innervation of the sclera. By contrast, stimuli applied to the iris initiated
high energy bursts of impulses, in keeping
with a rich innervation of epithelium and
vessels. Traction on the lens and pectinate
ligaments produced both high and low amplitude impulses. These may arise in receptors of the ciliary body, since lens and
zonule fibers are devoid of nerves. A proprioceptive function is inferred, though
differentiated terminations similar to somatic mechanoreceptors are absent.
Ocular sensation, a consideration in a
variety of clinical conditions, may be
aroused from both visceral and somatic
endings in the globe. In order of decreasing occurrence these are limited to vessels
of the iris, choroid, ciliary body and
limbus, and in cornea, adjacent conjunctiva, iridial epithelium, tendon insertions,
and sclera. Furious outbursts of high amplitude impulses were aroused by intraocular tension (Tower, ’40), even after
cornea and sclera were excluded. This degree of response could arise only from endings on vessels of choroid, iris, retina, and
ciliary body.
CONCLUSIONS
1. The innervation of the eye, excepting retina, was investigated. Our purpose
has been confirmation, extension, and correction of existing works.
2. Ciliary nerves constitute the sole nonvisual innervation of the globe. Their
motor fibers are destined for smooth muscle of the iris, ciliary body, and blood vessels. Sensory components are distributed
to the cornea, sclera, epithelium of ciliary
processes and anterior iris, and perhaps to
blood vessels. Sensory endings in the
choroid and ciliary body are probably
limited to visceral sensory functions, since
they occur only in relation to vessel walls.
3. Ganglion cells were not observed in
the course of ciliary nerves. If they exist
they are exceedingly rare, and are presumably displaced ciliary, sympathetic, or Gasserian ganglion cells.
4. “Trophic” endings in sclera (Agababow, ’12) were not found.
5. A terminal syncitium of axons in
either somatic or visceral structures, as observed by Boeke and Hirano, was nowhere
in evidence. All endings, whether sensory
or motor, were discrete.
.
INNERVATION OF EYE
LITERATURE CITED
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317-380.
Arnold, J . 1863 Uber die Nerven und das Epithelium der Iris. Virchows Arch., 27: 345-374.
Attias, G. 1912 Die Nerven der Hornhaut des
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Bach, L. 1896 Die Nerven in der Sclera mit
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Bietti, A. 1897 Sulla distributione della fibre
nervose del corpo ciliare. Ann. Ottal., 26: 215222.
1897 Le fibre nervose della coroidea
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Boeke, J. 1933 Innervationstudierum, 111. Ztschr.
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1935 Innervationstudierum, VIII. Ibid.,
38: 549-618.
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Crouch, R. 1936 The efferent fibers of the
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Dogiel, A. 1890 Die Nerven der Cornea des
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1891 Die Endkorperchen (Endkolben,
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bulbi des Menschen. Arch. Mikr. Anat., 37:
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Duke-Elder, W. 1932 Textbook of OphthalmoIogy, vol. I. Henry Kimpton, London.
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muscle. Anat. Rec., 127: 219-229.
Givner, I. 1939 Episcleral ganglion cells. Arch.
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Hirano, N. 1941 Nervose Innervation des Corpus ciliares des Menschen. v. Graefes Arch.
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Hosch, F. 1891 Erlichs Methyleneblau methode
und ihre Anwendung auf das Auge. Ibid., 37,
3: 37-54.
Hoyer, H. 1873 Uber die Nerven der Hornhaut.
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Kirpitschowa-Leontowitsch, W. 1911 Zur Frage
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Kiihne, M. 1862 Note sur nouvel ordere nerfs
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Lipmann, H. 1869 Uber die Endigung der
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Meyer, A. 1880 Die Nervenendigung in der Iris.
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Miller, M. 1960 The pattern of cutaneous innervation of the human hand, foot and breast.
In: Advances in Biology of Skin, vol. I. J. W.
Arrowsmith, Ltd., Bristol, England.
Pause, C. 1877 Uber die Nerven der Iris. V.
Graefes Arch. Ophthal., 23, 3: 1-23.
Ranvier, L. 1881 Lepons D’Anatomie Gbn6rale
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Ray, B., J. Hinsey and W. Geohegan 1943 Observations on the distribution of the sympathetic nerves to the pupil and upper extremity
as determined by stimulation of the anterior
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Schweigger, C. 1860 Uber die Ganglion Zellen
und blassen Nerven der Choroidea. v. Graefes
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Smirnow, A. 1900 Die weise Augenhaut als
stelle der Nerven endigungen. Anat. Anz., 18:
76-80.
Stohr, P. Jr. 1932 Mikroscopische Studieren zur
Innervation des Magen-Darmkanales, 11. Uber
die Nerven das menschlichen Magens und ihre
Veranderungen beim Ulkus. Ztsch. Zellforsch.,
16: 123-197.
Tower, S. 1940 Unit for sensory reception in
cornea: with notes on nerve impulses from
sclera, iris, and lens. J. Neurophysiol., 3: 486500.
Virchow, H. 1910 Mikroskopische Anatomie
der Auzeren Augenhaut und des Lidapparates.
Graeff-Saemisch Handbuch der Augenheilkunde,
2nd Aufl. Bd. I, Abt. I, K. 11, 183-229.
Waldeyer, W.,and V. Izquierdo 1880 uber die
Endigungweise der sensiblen Nerven. Arch.
Mikr. Anat., 17: 367-382.
Wolff, E. 1948 The Anatomy of the Eye and
Orbit, 3rd ed. The Blakiston Co. Philadelphia.
Zander, E., and G. Weddell 1951 Observations on the innervation of the cornea. J. Anat.,
85: 68-99.
INNERVATION OF EYE
J. C. Beatie and Donald L. Stilwell, Jr.
PLATE 1
1
Internal aspect of hemisphere of eye. The retina and the right quadrant of ciliary body and iris have been removed to show the course
of ciliary nerves which lie external to these structures. The numbers refer to photographic figures in plates 2-4. For clarity the extent
of ciliary nerve branchings, complexity of ciliary body and iridial arcades, and abundance of corneal innervation are simplified. The pericorneal plexus is indicated by 14, level of ciliary body by 9.
OPTIC
NERVE
PLATE 2
EXPLANATION OF FIGURES
Vascuhture, choroid, and sclem. These and all succeeding figures are photographs of
full-thickness, cleared preparations of eyes, stained supravitally with methylene blue.
2
Long ciliary nerve (right) and innervation of long posterior ciliary artery. Smaller paraarterial nervi vasorum parallel to course of vessel. Periarterial nerves form a plexus
around the vessel. X 150.
3 Para-arterial nerves divide to follow choroidal branches of long posterior ciliary artery.
Fat cells in lower center may be mistaken for ganglion cells. X 75.
54
4
Choroidal vein innervation, derived from nerves in figures 2 and 3.
5
Two free endings on internal surface of sclera. X 300.
x
300.
INNERVATION OF 'ZYE
J. C. Beatie and Dorald L. Stilwell, Jr.
PLATE 2
PLATE 3
9 Terminal innervation of ciliary body smooth muscle. Nerve fibers are parallel to muscle
bundles. x 250.
Portion of arcade nerve (bottom) and several tcrtuous, radially distributed branches in
iris. x 200.
200.
8
x
Innervation of predominantly circularly-oriented vessels in body of iris. X 300.
Plexiform distribution and termination of sensory nerves of iridial epithelium.
6
7
Iris and ciliary body
EXPLANATION OF FIGURES
INNERVATION OF EYE
J. C. Beatie and Donald L. Stilwell, Jr.
PLATE 3
Two large ciliary nerves in center of photograph send branches into cornea (above top
margin) to terminate in somatic sensory endings, seen in figures 12 and 13. Other
branches are directed laterally (circumferentially) to participate in the formation of
the pericorneal plexus. The plexus is seen as a network of smaller bundles across the
top one-third of this photograph. Plexus also contributed to by one of 40-50 small ciliary
nerve bundles (arrow). X 100. Terminal destinations of pericorneal plexus: ( a ) free
endings in peripheral one-third of cornea. See also figure 15. ( b ) Recurrent autonomic
fibers to nerve arcades in ciliary body (labeled); ( c ) recurrent free endings in fascia of
extra-ocular muscle insertions. See figure 11. ( d ) Autonomic innervation of vessels in
limbic area, and profuse free endings in epithelium of limbus (fig. 1 4 ) .
Recurrent fibers of pericorneal plexus, ending freely in fascia of rectus muscle insertion.
Above, pericorneal plexus; below, muscle attachment. Nerve at arrow is one of 40-50
smaller ciliary nerve bundles which contributes to plexus, similar to the one pointed out
i n figure 10. X 75.
10
11
Pericorneal plexus (limbic region)
EXPLANATION O F FIGURES
PLATE 4
INNERVATION OF EYE
J. C. Beatie and Donald L. Stilwell, Jr.
PLATE 4
Free endings in cornea. No terminal fusion of nerve elements is observed.
x
200.
15
Free ending at periphery of cornea, derived from a branch of pericorneal plexus.
x
300.
14 Free sensory and vasomotor fibers near surface of limbus, overlying elements of the
pericorneal plexus. Note close resemblance of this vascular innervation with that in
iris, figure 6. x 300.
12, 13
Cornea and limbus
EXPLANATION OF FIGURES
PLATE 5
i
h
61
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