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Structure and innervation of the ciliary processes of the albino rabbit eye.

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Structure and Innervation of the Ciliary
Processes of the Albino Rabbit E y e '
Department of Anatomy, Pennsylvania State University, The Milton S .
Hershey Medical Center, Hershey, Pennsylvania 17033
Various physiological and anatomical studies have suggested that the
ciliary epithelium may be under the influence of adrenergic nerve fibers. This study
was undertaken i n a n attempt to characterize the innervation of the ciliary epithelium.
Both light and electron microscopy revealed that the ciliary epithelium is only
sparsely innervated. In addition, the ciliary epithelium overlies a highly vascular
stroma, and the distance between the large vascular channels and the ciliary epithelium is small. These features of the stroma indicate it is structurally suited
for a transport function, a fact which supports earlier reports that the elaborate
membrane infoldings of the ciliary epithelium are indicative of a transport function.
The discrepancy between previous descriptions of the innervation of the ciliary epithelium, demonstrating complex subepithelial plexuses of adrenergic nerve fibers,
and the results of the present study may be due to the fact that the fluorescence
technique used i n previous studies could have labelled some other tissue component
i n addition to adrenergic nerve fibers. Results with orcein and aldehyde fuchsin
stains indicate that the disposition of elastic fibers in the ciliary processes is similar
to previous descriptions of adrenergic nerve fibers as determined by fluorescent
The ciliary body is designated as that
portion of the tunic coats of the eye between the anterior limit of the retina and
the base of the iris. It is generally divided
into two parts: the posterior pars plana, a
flattened area, and the anterior pars plicata, where the tunic coats are thrown into
a series of folds called ciliary processes.
These ciliary processes are presumed to be
the site of elaboration of aqueous humor,
the fluid responsible for maintaining intraocular pressure, and hence, the internal
integrity of the eye.
There is an abundance of literature
which indicates a relationship between the
autonomic nervous system, the rate of formation of aqueous humor, and intraocular
pressure. Various chemicals and drugs,
including several autonomic drugs, have a
significant effect on intraocular pressure
and the rate of aqueous humor formation.
These include pilocarpine (Edwards, Hallman and Perkins, '67), the carbonic anhydrase inhibitor acetazolamide (Tonney, '63;
Oppelt, '67), ethanol (Houle and Grand,
'67), ascorbic acid (Linner, '65), p-p'-iminodipropionitrile (Heath and Wang, '68),
norepinephrine (Hoffman, '68), and ouabain (Oppelt and White, '68; Waitzman
and Jackson, '65). It has also been demANAT.
REC.,168: 339-350.
onstrated that an increase in intraocular
pressure causes a decrease in the rate of
aqueous humor formation, and vice versa
(Bill, '67; Macri, '67). In addition, superior
cervical ganglionectomy results in reduction of intraocular pressure, an effect attributed to the release of stored catecholamines
in the degenerating postganglionic terminals (Crombie and Hendley, '67; Hendley
and Crombie, '67). Finally, monoamine
oxidase (MAO) is present in significant
quantity in the nonpigmented cell layer of
the ciliary epithelium (Shanthaveerappa
and Bourne, '64; Lukas and Cech, '65). The
function of MA0 in catecholamine metabolism sugggests that the ciliary epithelium may be under the influence of
adrenergic nerve fibers.
Despite some convincing evidence that
the autonomic nervous system may play
a role in aqueous humor formation, little
is known concerning the innervation of
ciliary epithelium. The present study was
undertaken in an attempt to define and
Received Jan. 5, '70. Accepted May 29, '70.
1 This research was supported in part by United
States Public Health Service Research grant AM 11407.
% T h i sresearch was carried out in part while the
investigator was the recipient of a summer research
fellowship awarded under United States Public Health
Service GRS grant 1 SO1 FR05680-01.
characterize the innervation of ciliary scopic study were exclusively from the long
anterior processes (Wegner, '67; Kozart,
Ultrastructural studies of ciliary epi- '68), and the base of the ciliary body was
thelium (Duke-Elder and Wybar, '61 ; removed by trimming the block. These proHolmberg, '59; Fine and Zimmerman, '63; cedures were followed due to the fact that
Tormey, '63, '65, '66) have clarified the large bundles of nerves can be found in
nature of the bilayer of epithelial cells, but the stroma beneath the ciliary body, courshave not commented on the nature of in- ing to the iris. Thin sections were cut on
nervation of the ciliary processes. Our a Porter-Blum MT-2 microtome and placed
knowledge concerning the innervation of on uncoated or formvar-coated grids. Secthis region of the eye is limited principally tions were stained with uranyl acetate and
to ciliary muscle (responsible for lens ac- lead hydroxide or with lead citrate, and
commodation) (Cogan, '37), pupillary examined with an RCA EMU-4 electron
muscles (Richardson, '64), and the drain- microscope.
age angle of the eye (Barany, '65).
Ehinger ('64, '66a,b, '67), using fluoresLight
In the sections
cence techniques for adrenergic nerve fibers, and Lukas and hIazanec ('67), in stained with hematoxylin and eosin, a
an electron microscopic study, have de- relatively basophilic bilayer of epithelium
scribed the presence of nerves near ciliary can be seen covering a highly vascular
epithelium, but the relationship of these stroma. Much of the stroma consists of
nerves to the epithelial cells of the ciliary large vascular channels; the remainder is
pale eosinophilic connective tissue.
processes cannot be evaluated precisely.
The sections stained with silver (fig. 1)
contain scattered, delicate nerve fibers loLight microscopy. The eyes of two al- cated predominantly in the thin connective
bino rabbits were removed, and longitudi- tissue, in close approximation to both the
nal sections containing the iris and ciliary vascular channels and pigmented epithelial
body were fixed in formalin for 24 hours. cell layer. These scattered fibers course to
The fixed sections were then cut into the very tip of a ciliary process. A few
wedge-shaped sectors and embedded in nerve fibers are closely applied to the epiparaffin. Sections 8 EL thick were stained thelium. Occasional nerve fibers are also
with hematoxylin and eosin, aldehyde located more centrally in the stroma of
fuchsin (Gomori, '50), and orcein (Pinkus, the ciliary processes in relationship to
'44). Sections 10 thick were stained by vascular channels. Both epithelial cell laythe silver method of Sevier and Munger ers stain a golden brown color.
The elastic tissue component of the
Electron microscopy. The eyes of five connective tissue was studied in sections
anesthetized albino rabbits were removed, stained with acid orcein-Giemsa as well
and longitudinal sections containing the as aldehyde fuchsin (fig. 2). Both elastica
iris and ciliary body were fixed in a for- stains demonstrate that plexiform arrangemaldehyde-glutaraldehyde solution and ments of elastic fibers are present within
postfixed in Oso,, according to the method the narrow connective tissue band beof Karnovsky ('65). Other sections of tween the vascular channels and the pigsimilar origin were fixed in a saturated mented epithelial cell layer. Elastic fibers
cold aqueous solution of KMnO,, accord- also course along the vascular channels,
ing to the method of Richardson ('66, '68). and near the apical portion of the nonpigAll sections were embedded flat in Araldite mented epithelial cell layer. Aldehyde
(Durcupan-Fluka). The embedded tissue fuchsin also stains zonular fibers, which
was oriented in vise-type holders so that take origin from the ciliary processes, the
the knife cut was in radial plane with deep purple color of elastica. The remainrespect to the orbit. This plane of section der of the connective tissue is a light bluepermitted precise phase microscopic locali- green color, representing fast green-positive
zation of the ciliary body in 1-2
thick collagen fibers. Epithelial cytoplasm stains
sections. The sections for electron micro- light red.
Fig. 1 Photomicrograph of the tip of a ciliary process stained with silver. Occasional
nerve fibers (arrows) are present beneath the bilayer of epithelium, as well as adjacent to
vascular channels. ( x 1000).
Fig. 2 Photomicrograph of a region of a ciliary process stained with aldehyde fuchsin.
Plexuses of elastic fibers are present beneath the bilayer of epithelium; the extensive distribution of elastica in the connective tissue stroma may also be seen. The heavy black lines
a t the apical borders of the process dcrnonstrate the elastica content of the zonular fibers.
Also apparent in this micrograph is the large size and frequency of vascular channels in a
ciliary process. ( X 1000).
Electron microscopy. The stroma underlying the epithelium consists of large
vascular channels i n a loose connective
tissue matrix (figs. 3 , 4 ) . The vascular
channels are quite large, often reaching a
size of 25 X 50 p. Because of the large
size and frequency of vascular channels
i n ciliary processes, when one surveys a
longitudinal section of a ciliary process
mounted on a grid, a vessel or part of a
vessel is present in almost every grid
square. The endothelium is thin, with occasional fenestrations and a prominent
basal lamina.
The connective tissue stroma between
the vascular channels and epithelium is
usually 1-2
thick, but occasionally expands to 6 p in width. The fibroblasts usually have long, thin cytoplasmic processes, and profiles of fibroblast cytoplasm
are scattered throughout the Connective
tissue. The fibroblast processes sometimes
extend almost to the basal lamina of the
epithelial cell layer. Collagen fibers and
occasional clumps of elastica are distributed throughout the connective tissue.
Nerve fibers and axonic expansions,
which could possibly represent terminals,
are only infrequently encountered. Fibers
devoid of associated Schwann cell cytoplasm were not observed. Axonic expansions in glutaraldehyde-fixed tissue (fig.
5, 6 ) contain many synaptic vesicles, most
of which are ungranulated. Some large
granulated vesicles and many mitochondria are also present in these regions of
the fiber. The axonic expansions are USually located 1 from the vascular channels and 4-5
from the base of the
pigmented epithelial cell layer. The expansions average 1.5 x 2.5 in size.
Axonic expansions in permanganatefixed tissue (fig. 7) are filled with synaptic
vesicles, the majority of which are the
small, granulated type. Occasional axonic
expansions containing ungranulated vesicles can also be seen. Usually a single
large mitochondrion is present in the
Nerve fibers (fig. 8) contain scattered
vesicles and few mitochondria. The fibers
lack the vesicle-laden appearance of the
axonic expansions described above. Nerve
fibers are usually 1 ,u. in diameter.
The fine structure of the two layers of
ciliary epithelium has been described previously (Tormey, '63, '65, '66).
The ciliary epithelium overlies a highly
vascular and sparsely innervated connective tissue stroma. This stroma is characterized by large vascular channels and
scant connective tissue between the vascular channels and the pigmented epithelial cell layer. As described previously
(Tormey, '63, '65, '66), the epithelial layers
have extensive membrane infoldings, especially along their lateral borders. The
epithelial cells also contain numerous
There is general agreement that the
ciliary processes are the site of aqueous
humor formation (Pappas, Smelser and
Brandt, '59; Duke-Elder and Wybar, '61;
Simon, Bonting and Hawkins, '62; Ham,
'65; Tormey, '65, '66; Duke-Elder and
Gloster, '68). The most widely accepted
theory of aqueous humor formation is that
i t involves both active transport and diffusion of plasma components from the
vascular channels of the ciliary processes
to the posterior chamber of the eye (DukeElder and Gloster, '68). Tormey ('65, '66)
concludes that the elaborate membrane
infoldings and high mitochondria1 content
of the ciliary epithelium strongly suggest
that the ciliary epithelium is a transport
epithelium. The present study would suggest that the connective tissue stroma of
the ciliary processes is also structurally
suited for a transport function. The vascular channels present in the ciliary processes are larger than would be expected if
these vessels functioned only to nourish
the area. The thin endothelium and numerous fenestrations are similar to those
in capillaries of renal glomeruli and most
endocrine glands. Fawcett ('63) states that
such a thin fenestrated endothelium facilitates diffusion and transport of plasma
components in to neighboring tissue. Finally, the short distance (1-2 p ) between
the vascular channels and the epithelial
cell layers would provide a minimum of
obstruction for the diffusion and transport
of substances.
With reference to the innervation of
ciliary processes, several discrepancies are
obvious between the results of the present
study and the previous studies of Ehinger
('66, '67) and Lukas and Mazanec ('67).
Using fluorescence techniques for adrenergic nerve fibers, Ehinger described a plexus
of adrenergic nerves in the connective
tissue adjacent to the pigmented epithelial
cell layer in the rabbit, as well as in other
animals. Although silver-stained sections
from the present study did reveal some
subepithelial nerve fibers, they gave no evidence of subepithelial plexuses of the
magnitude described by Ehinger. While i t
is not possible to precisely quantitate the
difference between the number of nerve
fibers found in the present study as compared with the findings of Ehinger, ultrastructural results in the present study also
revealed that nerves are present quite infrequently -being in evidence in only
5-10% of survey pictures of the subepithelial area.
In order to explain this discrepancy,
this author must conclude that the fluorescence technique labelled some other tissue
Fig. 3 Electron micrograph of a representative
area of a ciliary process. The bilayer of ciliary
epithelium overlies a thin band of loose connective tissue. A large vascular channel (VC), with
its thin, fenestrated endothelium, is present at
the lower right. One entire fibroblast, the cytoplasmic process of another, and diffuse collagen
fibers are present i n the connective tissue. The apical (nonpigmented) layer of epithelial cells (NPE)
is characterized by elaborate membrane infoldings. (Formaldehyde-glutaraldehyde. x 10,250).
Figure 3
Figure 4
Fig. 5A. Axonic expansion ( T ) located between the vascular channel and the basal
layer of ciliary epithelial cells. The thinness and occasional fenestrations (arrow) of the
endothelium are apparent i n this micrograph. (Formaldehyde-glutaraldehyde. x 10.250).
Fig. 5B The inset is a higher magnification of a region of figure 5A. The axonic expansion contains predominantly large ungranulated vesicles, with a few granulated vesicles.
Schwann cell cytoplasm partially surrounds the expansion. (Formaldehyde-glutaraldehyde.
X 45,200).
Fig. 4 A representative view of a ciliary process, fixed in permanganate. A large vascular
channel ( V C ) containing several distorted erythrocytes i s present in the lower center of the
micrograph. A prominent endothelial basement
membrane encircles the vascular channel. Several
nerve fibers (arrows) associated with Schwann
cell cytoplasm appear i n the thin connective
tissue between the vascular channel and the
epithelium. Large mitochondria ( M ) , swollen by
permanganate, and characteristic membrane infoldings are present in the bilayer of ciliary
epithelium at upper left and center of the micrograph. x 8000).
component in addition to nerve fibers.
Aldehyde fuchsin and orcein-stained sections displayed a plexiform arrangement
of elastica in the subepithelial connective
tissue, and intimately investing the vascular channels (fig. 2). It would seem that
this disposition of elastica is remarkably
similar to the organization and location of
Ehinger’s adrenergic nerve fibers. To determine histochemically whether or not
elastica and adrenergic nerve fibers react
similarly to the fluorescence technique
Fig. 6 Large axonic expansion ( T ) . Numerous ungranulated vesicles and mitochondria are present i n the expansion, which is partially enclosed by Schwann cell cytoplasm. The junction of two
endothelial cells can be seen below the Schwann cell. (Formaldehyde-glutaraldehyde. x 34,200).
Fig. 7 Axonic expansions. Several small, granulated vesicles and a single large mitochondrion
appear in the larger expansion. Cytoplasm of a single Schwann cell partially surrounds both expansions. The basal lamina of a pigmented epithelial cell is present i n the upper left corner of micrograph. (Permanganate. x 26,100).
Fig. 8 Bundle of nerve fibers. Several adrenergic fibers are present, as well as a fiber (arrow)
which lacks the typical appearance of an adrenergic nerve fiber. Part of a pigmented epithelial cell appears i n the upper left corner of the micrograph. (Permanganate. x 19.500).
used by Ehinger is quite outside the scope
of the present study. Perhaps this question
will be more easily answered when the
exact composition of elastica is determined. Nevertheless, in light of the similarity between the organization of elastic
fibers in the present study and the disposition of adrenergic nerve fibers as described
by Ehinger, one cannot deny the possibility
that elastica does indeed react to the
fluorescence technique of Ehinger.
Lukas and Mazanec ('67) report that
large bundles of adrenergic axons are
present close to the ciliary epithelium. The
results of the present study would indicate
that large bundles of axons are not present
in the ciliary processes. It seems likely that
Lukas and Mazanec encountered nerves
in the ciliary body, perhaps at the base
of ciliary processes, but not in ciliary
processes themselves.
Richardson ('66, '68, '69) has developed
a technique involving permanganate fixation which distinguishes between adrenergic and cholinergic nerve fibers, and he has
described the criteria upon which such a
distinction can be made. On the basis of
Kichardson's criteria, the results of this
investigation confirm the presence of adrenergic nerve fibers in ciliary processes.
The finding of adrenergic nerve fibers, with
their typical small, granulated vesicles, was
expected. There are many reports in the
literature which suggest an adrenergic innervation of the ciliary processes. Electrical stimulation of the cervical symDathetic
nerve (Langham and Rosenthal, '66), superior cervical ganglionectomy (Langham,
'65; Crombie and Hendley, '67; Hendley
and Crombie, '67) and pharmacologic
( guanethidine-induced)
(Anselmi, Bron and Maurice, '68) all
caused significant decrease of intraocular
pressure and aqueous humor formation
rate. Although the possibility exists that
these effects are secondary to changes in
blood flow in the ciliary processes, i t seems
more likely that sympathetic nerves have
some direct effect on aqueous humor secretion. Langham and Rosenthal ('66)
suggest the possibility of a direct inhibitory
effect of the sympathetic transmitter on
aqueous humor formation. This suggestion
is supported by the finding in the present
study that nerve fibers and terminals are
close to both vascular channels of the
ciliary processes and epithelial cell layers.
Langham ('65) speculates that two sympathetic mechanisms may be involved in
the effects described above : a-adrenergic
fibers causing changes in the outflow facility of the eye, and p-adrenergic fibers in
the ciliary processes which inhibit aqueous
humor secretion.
In the present study, this author occasionally encountered nerve fibers which
lacked the typical appearance of adrenergic fibers. These infrequent fibers were
characterized by a paucity of vesicles, most
of which were ungranulated. Although
these may in fact be adrenergic fibers, one
cannot exclude the possibility that these
fibers may be cholinergic. Indeed, Edwards
et al. (Edwards, Hallman and Perkins,
'67) report that administration of the
cholinergic drug, pilocarpine, caused a significant increase in the rate of aqueous
humor formation. Further investigation is
needed to determine the exact nature of
these nerve fibers.
The author acknowledges the valuable
advice of Dr. Bryce L. Munger.
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albina, structure, rabbits, processes, ciliary, innervation, eye
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