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© 2004 American College of Veterinary Ophthalmologists

Veterinary Ophthalmology

(2004)

7

, 1, 11–22

Blackwell Publishing Ltd.

REVIEW

TAPETUMLUCIDUMAMONGVERTEBRATES

Comparative morphology of the tapetum lucidum (among selected
species)

F. J. Ollivier,* D. A. Samuelson, D. E. Brooks, P. A. Lewis, M. E. Kallberg and A. M. Komáromy

Department of Small Animal Clinical Sciences, College of Veterinary Medicine, University of Florida, Gainesville, Florida, USA

Abstract

Objectives

The phenomenon of ‘eye-shine’ is seen in a variety of animal species, and is
generally thought to be related to the presence of an intraocular reflecting structure, the
tapetum lucidum. The tapetum lucidum is a biologic reflector system that is a common
feature in the eyes of vertebrates. It normally functions to provide the light-sensitive
retinal cells with a second opportunity for photon-photoreceptor stimulation, thereby
enhancing visual sensitivity at low light levels. The tapetum lucidum is presented here
according to a classification based on the location, as well as the composition, of this
reflective layer. Finally, the physical and chemical properties, as well as the origins of the
different tapeta lucida, are discussed and compared.

Methods

The anatomic and biochemical aspects of the tapetum lucidum in various
vertebrates are examined. Morphologic observations were made from paraffin and plastic
embedded specimens. Specimens were treated with traditional stains and observed by
light and transmission electron microscopy.

Results

Some species (primates, squirrels, birds, red kangaroo and pig) do not have this
structure and they usually are diurnal animals. In vertebrates, the tapetum lucidum
exhibits diverse structure, organization and composition. Therefore, the retinal tapetum
(teleosts, crocodilians, marsupials, fruit bat), the choroidal guanine tapetum
(elasmobranchs), the choroidal tapetum cellulosum (carnivores, rodents, cetacea), and
the choroidal tapetum fibrosum (cow, sheep, goat, horse) are described.

Conclusions

The tapetum lucidum represents a remarkable example of neural cell and
tissue specialization as an adaptation to a dim light environment and, despite these
differences, all tapetal variants act to increase retinal sensitivity by reflecting light back
through the photoreceptor layer. These variations regarding both its location and
structure, as well as the choice of reflective material, may represent selective visual
adaptations associated with their feeding behavior, in response to the use of specific
wavelengths and amount of reflectance required.

Key Words:

morphology, tapetum lucidum, vertebrates

Address communications to:
Franck Ollivier
Tel.: (352) 392–4700
Fax: (352) 392–6125
e-mail:
ollivierf@mail.vetmed.ufl.edu
*Present address:
Department of Small Animal
Clinical Sciences
College of Veterinary Medicine
University of Florida
Gainesville, FL 32601–0126
USA

INTRODUCTION

The visual senses include perceptions of light, color, form,
space and movement. The light sense, by which light is
perceived as such, and gradations in its intensity appreciated,
is the most fundamental of the visual senses and it is highly
developed in vertebrates.

1

The attainment of a high standard
of light sensitivity involves certain structural specializations
in the eye. The eyes of animals to which an acute perception
of light is a necessity can be differentiated from animals which
need excellent visual acuity. From this point of view verte-
brates can be divided into three main classes: diurnal animals
(adapted to bright light), nocturnal animals (adapted to near-
darkness of night), and arrhythmic animals (adapted either
to bright or dim illumination). The ocular microanatomy
of a nocturnal and a diurnal eye are very different, with
compromises needed in the arrhythmic eye. Anatomic
differences in light gathering are found in the organization
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of the retina and the optical system. The size of the eye and
those of its different structures influence the amount of light
gathered. The presence of a tapetum lucidum influences the
light sensitivity capabilities of a species.
The phenomenon of ‘eye-shine’ is seen in a variety of animal
species, and is generally thought to be related to the presence
of an intraocular reflecting structure, the tapetum lucidum.

2,3

The tapetum lucidum is a biologic reflector system that is a
common feature in the eyes of vertebrates. It normally func-
tions at low light levels to provide the light-sensitive retinal
cells with a second opportunity for photon-photoreceptor
stimulation, thereby enhancing visual sensitivity (Fig. 1).

4,5

While
increasing retinal sensitivity in dim illumination, the tape-
tum lucidum may harm visual acuity in bright illumination.

4

Tapeta lucida in vertebrates exhibit diverse structure, organ-
ization and composition (Fig. 2). Despite these differences,
all tapetal variants act to increase retinal sensitivity by
reflecting light back through the photoreceptor layer. Tapeta
lucida can be classified according to the location as well as
the nature of the reflective materials that constitute them.
The reflecting material may be located within the cytoplasm
of the retinal epithelium (retinal tapetum lucidum) or in the
choroid. The choroidal tapetum lucidum lies immediately
adjacent to the choriocapillaris and external to the retinal
epithelium (Fig. 1). The reflective material of the choroidal
tapetum lucidum may be a guanine tapetum, an array of
extracellular fibers (tapetum lucidum fibrosum), or a layer of
cells (which compose the tapetum lucidum cellulosum) packed
with organized, highly refractive material (tapetal rodlets)
(Fig. 1).

3,5,6

This paper is a review of the anatomic, histologic and bio-
chemical aspects of the tapetum lucidum of various verte-
brates. Three basic tapetal morphotypes of vertebrates are
described and compared: the absence of tapetum, the retinal
tapetum lucidum, and the choroidal tapetum lucidum.

ABSENCE OF TAPETUM

Animals without a tapetum lucidum (e.g. primates, squir-
rels, birds, red kangaroo and pig) generally have diurnal
habits, and a red or orange to pale gray fundus reflection
(Fig. 2, lane 1).

7,8

The red to orange background results from
reflection of light from choroidal blood vessels.

2

Birds are
apparently the only large group of animals in which the tape-
tum is consistently absent.

16

RETINAL TAPETUM LUCIDUM

Retinal tapeta lucida are common in fish and some reptiles, but
rarely occur in mammals.

3,6,9

In most of these species the retinal
tapetum can be classified as either a lipid (in some teleosts, mam-
mals) or guanine type (in some teleosts, reptiles) (Table 1).
Another differentiation can be made among the retinal
tapeta lucida based on whether the tapetum is occlusible (found
in most of teleosts), or nonocclusible (or static) (found in
mammals). The variation in ‘eye-shine’ in occlusible tapeta
in teleosts is caused by the migration of black pigment (mel-
anin) within the retinal pigment epithelium (RPE) and the
choroidal cells. Melanin migrates towards the vitreous when
the eye is illuminated and towards the sclera (backwards) in
dim light or darkness (Fig. 1). The migration of melanin can
mask or unmask the reflecting material depending upon the
degree of environmental illumination.

10–12

In certain species the tapetum is occlusible not because of
a pigment migration within a cell, but because of a visual cell
movement called the retinomotor movement, as well as the
migration of the processes of the RPE cells between the cells
and over the reflective material (elasmobranchs).

12,13

It is thought that the function of the occlusible tapetum is
related primarily to reduction of ‘eye-shine’ rather than modify-
ing retinal sensitivity, as the tapetal reflection contributes only
30% of the light reaching the photoreceptors (70% of the
light directly reaches the photoreceptors).

14,19

Fish – teleosts

Two morphologic types of tapeta lucida can be observed in the
eyes of fishes: one that lies in the choroid such as occurs in
elasmobranchs (that will be discussed later); and the other
that lies in the pigment epithelium of the retina such as occurs
in teleosts.

1,11,14

The choroid of teleosts is a connective tissue layer rich in
blood vessels, and one or two choroidal glands. A choroidal
tapetum of reflecting cells lying immediately outside the
choriocapillaris is formed in a few primitive teleosts. More
commonly, the tapetum lucidum is an integral part of the
pigment epithelium in primitive teleosts (lying in the pro-
cesses of the pigmented epithelial cells) ( Fig. 3).

11,12

The reflec-
tive material is contained in spheres closely packed together
in the apical processes of the pigment epithelial cells. It can
vary in chemical nature (Table 1). The tapetum of the seatrout
(

Sciaenidae

) contains highly refractive lipid (triglycerides)
particles,

10,12

whereas the tapetum of gars (

Lepisosteidae

) is
composed of a yellow phenolic compound.

11

The lipid tape-
tum observed in the seatrout has a white appearance, and
reflects light diffusely, as the reflective material is not organ-
ized within spheres.

10,11

In contrast, the tapetum of gars pos-
sesses a yellow reflective material (yellow pigment) that
occurs in regular array and is contained in a myriad of small
spheres. The spheres are organized in parallel sheets with
packing being hexagonal in the plane of the sheet.

11

The ‘eye-
shine’ varies in color (from yellow to dark brown) depending
on the retinomotor movement on the stage of light adapta-
tion of the eye, and characterizes the tapetum of these teleosts
as occlusible (Fig. 1).

10–12

The stratum argenteum is a ‘silvery’ reflective layer in the
outer choroid, adjacent to the sclera, frequently present in
fish larvae but not as frequent in adults. In species known to
have a stratum argenteum as adults, like the toadfish (

Toad-
fishes

), the tapetum lucidum is absent or vestigial and there is
little choroidal melanin.

12

The choroid of toad fish is thin and contains blood vessels,
a choroidal gland, and a stratum argenteum. There is loose
© 2004 American College of Veterinary Ophthalmologists,

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connective tissue between the choriocapillaris and the stra-
tum argenteum. Melanocytes are absent except for a few
near the globe equator. The outermost layer of the choroid is
occupied by a thin stratum argenteum which lies adjacent to
the sclera, is bilaminate, and becomes a layer of pentameric
crystals and a layer of yellow birefringent granules. The
inner surface of the stratum is shiny, has an orange metallic
luster, and is mostly a specular reflector, reflecting much of
the incident light. It consists of two or more cell layers (iri-
dophores). The iridophores are polygonal in shape and
loaded with packets of elongated thin pentagonal crystals
that are oriented with their long axis parallel to the argenteal
surface. The thin crystals are superposed over each other
and form a good reflective system. The reflective material is
uric acid (Table 1). The outer surface of the argenteum is
covered by a thin yellow sheet that has a mat appearance.
The yellow packets are in the form of 1-

µ

m diameter small
Figure 4. The retinal tapetum lucidum in alligator (Alligator mississipiensis).
inl, inner nuclear layer; onl, outer nuclear layer; r & c, rods & cones;
t /rpe, tapetum/retinal pigment epithelium; ch, choroid; sc, sclera.
(H&E staining, original magnification ×200, insert ×400.)
Figure 5. The retinal tapetum lucidum of a fruit bat (Pteroptus giganteus).
inl, inner nuclear layer; onl, outer nuclear layer; r & c, rods & cones;
t /rpe, tapetum/retinal pigment epithelium; ch, choroid; sc, sclera.
(H&E staining, original magnification ×200.)
Figure 1. Diagram of the three types of the tapetum lucidum present
among vertebrates. 1. Incoming light ray; 2. First stimulation on the
rods and cones; 3. Reflection on the tapetum lucidum; 4. Second
stimulation on the rods and cones.
Figure 2. Ocular fundus of various species showing the absence or the
presence a tapetum lucidum that can exhibit various colors. From right
to left: lane 1: absence of tapetum: bird, albinos rat, hooded rat, red
kangaroo, pig, rhesus monkey, human. Lane 2: retinal tapeta: alligator,
possum, fruit bat. Lane 3: Choroidal tapeta cellulosa: puppy, dog (3), cat,
jaguar, leopard. Lane 4: choroidal tapeta fibrosa: foal, horse, sheep, goat,
cow, deer, mouflon.
Figure 3. The retinal tapetum lucidum in fish (teleosts): oscar
(Astronotus oscellatus). onl, outer nuclear layer; r & c, rods and cones;
t /rp, tapetum/retinal pigment epithelium; ch, choroid. (H&E staining,
original magnification ×400.)
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granules, appear to have a geometric form, and are packed
closely in a continuous mass. By moving posteriorly or ante-
riorly within the cells, both the choroidal and the retinal
pigment prevent the light from reaching the argenteum
(Fig. 1) and retina. Therefore, the tapetum is occlusible in
toadfish.

12

Reptiles – crocodilians

In crocodilians, a semicircular area of the retinal epithelium
in the superior fundus shows a fair sulfur yellow coloration and
is specialized as a tapetum lucidum (Fig. 2, lane 2, #1).

9,15

Guanine crystals or calcium salts of guanine appear to be the
most common reflective material in this tapetum lucidum
(Table 1).

9,15,16

Nearly all of the epithelial cells contain
membrane-bound tapetal crystals. The crystals measure
100–120 nm in thickness and up to 0.7

µ

m in length.

9,15

In
the

Caiman crocodilus

the guanine crystals are highly organ-
ized at a cellular level, being perpendicular to the long axis of
the retinal photoreceptors (e.g. to the path of light).

9

In con-
trast, no such regular orientation of the guanine has been
seen in the spectacled caiman (

Caiman sclerops

).

15

In specta-
cled caiman, the reflecting material is much closer to the
photoreceptors, which may allow dispersion of the reflected
light to occur more readily. In the tapetal area, no melanosomes
are formed indicating a nonocclusible tapetum in the

Caiman sclerops

.

15

At the edge of the tapetum, however, both
guanine crystals and melanosomes are found within the epi-
thelial cells.

15

In comparison, the dark coloration of the infe-
rior part of the fundus in the

Caiman crocodilus

is caused by
masking of tapetal crystals by melanosomes; therefore, the
tapetum is an occlusible type. Outer segments of the retinal
receptors are deeply inserted into broad, mainly crystal-
containing processes of the pigment epithelial cells.

9

Other
than the presence of guanine crystals, the fine structure of the
retinal pigment epithelial cells of the crocodilians in general
resembles that found in other vertebrates (Fig. 4).

15
Figure 6. The tapetum lucidum cellulosum consists
of layers tapetal cells regularly arranged. Tapetum
cellulosum of the dog (a), the mink (b), the cat
(c), and the cougar (d). inl, inner nuclear layer; onl,
outer nuclear layer; r & c, rods & cones; t, tapetum;
rpe, retinal pigment epithelium; ch, choroid; sc,
sclera; c, capillary. (Trichrome staining, original
magnification ×200, insert ×400.)
Figure 7. Electronic microscopy of the tapetal
area in a dog (a) and a cat (b). The rodlets in the
tapetal cells are more precisely oriented in the
tapetum of the cat than in the tapetum of the dog.
os, outer segment; rpe, retinal pigment epithelium;
t, tapetum; tc, tapetal cell; c, capillary. (Original
magnification ×4000.)
© 2004 American College of Veterinary Ophthalmologists,

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Mammals – marsupials – opossum

A tapetum lucidum is present in a semicircular area of the
superior retinal epithelium of the opossum (

Didelphis virgin-
iana

) (Fig. 2, lane 2, #2). The reflecting material consists of a
large quantity of lipoidal spheres scattered throughout the
epithelial cells. The size and shape of the lipoidal spheres
vary somewhat from cell to cell, but all of the spheres within
the same epithelial cell are generally of a similar morpho-
logic appearance.

1,16,17

The reflecting material contains cho-
lesterol or cholesterol esters (Table 1).

16,17

Centrally in the
tapetal area, very few or no melanosomes are found, indicat-
ing a nonocclusible tapetum. At the periphery, the epithelial
cells contain both reflecting material and melanosomes. The
choriocapillaris over the tapetal area is organized in a similar
way to other mammals, as will be discussed later.

17

Mammals – fruit bats

The tapetum lucidum in fruit bats (

Pteroptus giganteus

)
appears white to yellow when isolated

in vitro

(Fig. 2, lane 2,
#3), but the reflected color is red

in vivo

, probably due to ret-
inal blood. The tapetal material lies at the base of the retinal
cells (which are arranged in a zig-zag manner) on the inner
side of the cones of pigment (Fig. 5). The reflecting material
is believed to be contained in many spherules and to be
phospholipid in composition (Table 1).

1,11,13,16,18

CHOROIDAL TAPETUM LUCIDUM

Guanine tapetum
Fish – elasmobranchs

Two morphologic types of tapetum
lucidum can be observed in the eyes of fish: one in the choroid
that occurs in elasmobranchs, and the other which occurs in
teleosts and lies in the pigment epithelium of the retina.

1,11,14

The elasmobranch tapetum in

Squalus

and

Scyliorhinus

con-
sists of sheets of 12–13 superposed guanine crystals in the
choroid. The thickness of the crystals ranges from 60 to
80 nm, which corresponds to the measured reflectance peaks
around 470–520 nm (Table 2). This type of biologic photon
reflector system approximates ‘ideal’ multilayer reflectors,
although three features weaken its capability. The reflectance
Figure 8. The tapetum lucidum fibrosum consists
of layers of collagen fibers regularly arranged. Tapetum
fibrosum of the horse (a), the sheep (b), the dolphin
(c), and the pilot whale (d). inl, inner nuclear
layer; onl, outer nuclear layer; r & c, rods & cones; t,
tapetum; rpe, retinal pigment epithelium; ch,
choroid; sc, sclera; c, capillary. (Trichrome staining,
original magnification ×200, insert ×400.)
Figure 9. Melanocytes may be the progenitor cell of the tapetal cells.
A tapetal cell in the tapetum cellulosum of the mink containing melanin
granules (m). tc, tapetal cell; c, capillary; rpe, retinal pigment epithelium;
m, melanin granules. (Trichrome staining, original magnification ×1000.)
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peak is much too broad, the arrangement of the reflective surface
is not always parallel to the retinal plane, and the presence of
migrating retinal epithelial cells in

Squalus

(an occlusible
tapetum) results in optical interference. The elasmobranchs
that have a choroidal tapeta often have very elaborate
occlusible tapeta lucida, as in some teleosts. The pigment in
retinal epithelial cells overlying the tapetum of the

Squalus

can migrate between and over the reflective cells during
illumination, and retreat during darkness to expose the reflectors
(Fig. 1). Retinal epithelial pigment migration may also occur
to limit exposure of an animal with ‘eye-shine’, as light in the
sea is very directional, and ‘eye-shine’ might increase the
chance of being caught by predators.
The choriodal tapetum of the Port Jackson shark (

Hetro-
dontus phillipi

) consists of a single layer of overlapping cells
situated immediately external to the choriocapillaris, and is
oriented at an angle of 30

°

to incoming light.

20,21

These tapetal
cells alternate with, and are separated from one another by
melanocytes that extend beyond the tapetal cells to intervene
between the tapetal cells and the incoming light. The bulk of
the tapetal cells are packed with evenly spaced guanine crystals.
The size and spacing of these reflective crystals is commensurate
with the principles of constructive interference. The crystals
measure approximately 0.1

µ

m in width and 5.0–8.0

µ

m in
length, and are separated by a space of 0.25

µ

m (Table 2).
These values are very similar to those found in other elasmo-
branchs (short-tailed stingray and southern fiddler ray

20,22

).
A typical tapetal cell will have 15–20 layers of these reflective
crystals throughout the thickness of the cell. The retinal
tapetum of the Port Jackson shark is an occlusible type.

Tapetum lucidum cellulosum
Mammals – carnivores
Common features

Many mammalian carnivores have a
choroidal tapetum of a cellular type (Table 2).

2

Histologically,
the tapetum cellulosum is composed of layers of rectangular-
shaped cells ( Fig. 1). The choroidal stroma adjacent to the
tapetum lucidum contains numerous small blood vessels that
penetrate the tapetal layer to form a single-layered capillary
network, known as the choriocapillaris, on the surface of the
tapetum. The tapetum cellulosum is thus interposed between
the large branching vessels in the choroid and the single
layer of the choriocapillaris beneath the retinal epithelium.
The thickness of the tapetum varies, being multilayered
at its center, and thinning to a single cell (or lamella) at its
periphery and adjacent to the optic nerve.

3,6,8,23–26

The tapetal cells are gradually replaced by melanin-
containing cells of the choroid at the outer edge of the tape-
tum (Fig. 1). At the periphery of the tapetum melanocytes
often intermingle with tapetal cells, and it is not uncommon
to observe cells which display morphologic characteristics of
both tapetal cells and melanocytes.

5,25

The layers of melano-
cytes gradually increase toward the periphery of the tape-
tum.

8

Over the tapetal region, the retinal epithelium (RE) is
nonpigmented to allow light to pass to and from the reflective
tapetum. At the periphery of the tapetum the RE becomes
pigmented. In nontapetal locations the RE is heavily pig-
mented to become the retinal pigment epithelium (RPE).
Histologically, the tapetal melanocytes are as thoroughly
dominated by a single type of cytoplasmic component (mel-
anin granules) as the tapetal cells. The principal difference is
in the high degree of order of the rods in the tapetal cells
compared to the random packing of the melanin granules in
the melanocytes.
The tapetum is supplied by branches of blood vessels of
the pigmented choroid. These penetrating vessels run from
the outer vascular layer of the choroid, pass through the
tapetum lucidum at right angles and terminate in the exten-
sive, anastomosing, capillary network: the choriocapillaris.
These capillaries form a single layer which underlie all parts
of the retina, are heavily fenestrated and are separated from
the tapetum or pigment cells of the choroid by a narrow
Table 1. Retinal tapeta among the vertebrates
Classification Type of tapetum Reflective material (nature, shape, size)
Fishes
Teleosts
Seatrout
10,11
Occlusible Spherules triglycerides; diameter: 0.4 µm
Gars
11
Occlusible Spherules yellow pigment (Phenolic compound); diameter: 0.24 µm
Toadfish
12
Argenteum stratum Acid uric
Occlusible
Reptiles
Crocodiles
Alligator mississipiensis
13
Spherules guanine
Caiman crocodilus
9
Occlusible Spherules guanine; diameter: 0.7 µm; length: 0.12 µm
Caiman sclerops
15
Non-occlusible Guanine
Marsupials
Opossum
16,17
Non-occlusible Lipoidal spherules; cholesterol and esters˙
Placentals
Bats
Fruit bat (Eidelon)
13,16
Spherules
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Table 2. Choroidal tapeta among the vertebrates


Classification Type of tapetum Reflective material
Thickness;
number of cell layers or crystals
Size of fibrils or
Rodlet’s dimensions
Fishes
Elasmobranchs, Occlusible Guanine crystals 60–80 µm 12–13 superposed crystals
Squalus
13,14,19
Elasmobranchs, Occlusible Guanine Length: 5–8 µm 15–20 superposed crystals
Port Jackson Shark
13,20,21
Width: 0.1 µm
Placentals
Carnivores
Dog
8,27
Cellulosum Zinc cysteine 9–15 layers Diameter: 4 µm
Length: 0.14 µm
Spacing: 0.20 µm
Zinc cysteine 9–11 layers
26
15–20 layers
3
Diameter: 4 µm
Length: 0.18 µm
Spacing: 0.20 µm
43 µm; 10–12 layers
41
Cat
5,18,25
Cellulosum Riboflavin 15–20 layers Diameter: 4–6 µm
Length: 0.10–0.12 µm
Spacing: 0.15 µm
Riboflavin, Zinc 16–20 layers
26
30 layers
8
18–30 layers
23
Diameter: 8–10 µm
Length: 0.15 µm
Spacing: 0.08 µm
6–10 layers
40
Diameter: 5 µm
Length: 0.23 µm
Spacing: 0.40 µm
35 layers
6
Ferret
26
Cellulosum Zinc cysteine 7–10 layers
Zinc cysteine 5–7 layers
30
Diameter: 3–4 µm
Length: 0.13–0.15 µm
50 µm; 10–15 layers
27
Diameter: 6 µm
Length: 0.20 µm
Spacing: 0.30 µm
Ranch Mink
31
Cellulosum 30 µm; 2–3 layers Non regular
Pinnipedes
Seal (Grey Seal)
4
Cellulosum ? 30–35 layers Diameter: 5–6 µm
Length: 0.10 µm
Spacing: 0.15 µm
Ungulates
Cow
36
Fibrosum Diameter: 0.15 µm
Spacing: 0.20 µm
Diameter: 0.20 µm
16
Sheep
38
Fibrosum Collagen 100–200 µm Diameter: 0.15 µm
Spacing: 0.20 µm
100–200 µm
23
Diameter: 0.15 µm
Spacing: 0.08 µm
140 rows per layer: 9–16 layers
26
Diameter: 0.15 µm
Spacing: 0.18 µm
Lemurs
Lemur (Galago Riboflavin
crassicaudatus)
16
Rodents
Paca (Cuniculus paca)
32
Fibrosum ? 12–15 Diameter: 8–10 µm
Length: 0.2–0.25 µm
Spacing: 0.15 µm
Primates
Non human (Aotus)
16
Fibrosum Collagen
18 or r i v i r r E T A L .
© 2004 American College of Veterinary Ophthalmologists, Veterinary Ophthalmology, 7, 11–22
layer of connective tissue that extends over the retinal sur-
face. In the tapetal and peritapetal areas, the choriocapillaris
is indented into the retinal epithelial layer leaving a flat inner
surface to the reflecting tapetum, while in peripheral regions
of the nontapetal fundus the RE is not indented by the
choriocapillaris.
3,5,6,25–27
The tapetum lucidum develops late in the eyes of some
animals (e.g. dogs and cats), and is usually completely devel-
oped by 4 months of age (Fig. 2, lane 3, #1). Animals born with
mature eyes have a well-developed tapetum although some
tapetal maturation occurs after birth. The degree of pigmen-
tation of the animal apparently has no effect on development
of the tapetum, with the exception of individuals with partial
or full albinism.
Cat ( Felis catus) The tapetum lucidum in the cat covers
nearly 50% of the fundus area, and is shaped roughly like a
rounded equilateral triangle, with its apex upwards and its
base passing horizontally just below the optic disc ( Fig. 2, lane
3, #5). Tapetal development initially produces the immature
feline tapetum that is pale blue in color ( Fig. 2). Its predom-
inant adult color varies from yellow to green between indi-
viduals (Fig. 2). Though the tapetum is undetectable grossly
in the new-born, it becomes obvious in 3- to 4-week-old kit-
tens, having essentially completed development at this time.
Subsequent enlargement of the eye causes the relative extent
of the tapetum to be reduced.
5,6,24,25
Histologically, the feline tapetum consists of 15–20 layers
of cells in its center, progressively thins and eventually disap-
pears towards the periphery (Table 2, Fig. 6c).
5,6,8,23–25
Aging
results in a loss of tapetal layers such that the adult cat tapetum
has fewer layers than the kitten.
5,25,26
In sagittal planes the individual feline tapetal cells appear
rectangular, set in even brick-like rows, and show an orderly
arrangement (Fig. 6c).
6
However, feline tapetal cells (located
close to the retinal epithelial layer) are flattened (thickness of
3–4 µm), having the largest surface parallel to the retina.
Only a small part of the tapetal cell is occupied by an ovoid
and centrally located nucleus. The bulk of the cell is filled
with numerous, electron-dense, membrane-bound rodlets of
reflective riboflavin (Fig. 7b).
24,25,28
The rodlets are 4–6 µm
in length and 0.1–0.12 µm in diameter, and are precisely ar-
ranged in a hexagonal lattice pattern with a space of 0.15 µm
separating adjacent rodlets (Table 2).
25
These rodlets are
grouped in bundles, which normally extend through the
entire thickness of the cell. Within each bundle the centers
of the rodlets are arranged parallel to the retinal surface with
remarkable regularity (Fig. 7b).
24
At the border region of the
tapetum, the feline tapetal cells sometimes contain melanin
granules and, similarly, some tapetal cell-like melanocytes
contain tapetal rodlets.
8,29
Dog (Canis familiaris) The tapetum lucidum in the dog is a
rounded equilateral triangle in shape, with the base oriented
horizontally (excluding the optic disc in most of the breeds)
and the apex directed upwards.
3,26
It covers an area of
approximately 30% of the superior fundus. In fresh prepara-
tions the reflected color of the canine tapetum varies from
yellow-green to green-blue with an irregular marginal area
(Fig. 2, lane 3, #2–4). The tapetum in the center of the tapetal
fundus of the dog consists of 18–20 layers of cells (Table 2),
with each layer containing approximately 15–18 ranks of
rodlets. The number of cell layers gradually thins towards
the periphery where it ends.
3,26
Tapetal cells appear as polyg-
onal plates arranged much like ‘patio bricks’ within each
layer. Sagittally, the canine tapetum resembles a multiple
layered ‘brick wall’ (Fig. 6a). Each cell is a rounded polygon,
40–48 µm in diameter and an average thickness of 3.28 µm.
The cytoplasm is tightly packed with separate bundles of
membrane-bound rodlets that displace other organelles
and/or inclusions into the perinuclear or peripheral region
(Fig. 7a).
3
Individual tapetal rodlets are fairly constant in
their dimensions with a mean length of 4.0 µm and a diame-
ter of 0.14 µm, and appear to contain a hard and apparently
homogeneous core (Table 2). Within the bundle rodlets are
primarily arranged into a compact regular hexagonal array,
with a mean spacing of 0.2 µm (Table 2). In the dog, Bruch’s
membrane over tapetal areas is extremely attenuated, appear-
ing as a single homogeneous layer (although it is bilaminate),
with a mean thickness of 0.16 µm.
Ferret (Mustela putorius) The tapetum lucidum in the fer-
ret is an iridescent triangular area in the rear pole of the eye
above the optic disc. It is present in both the pigmented and
albino eye.
27,30
A comparison of pigmented and albino fer-
rets revealed no morphologic differences of their cellular
tapeta at both light microscopic and transmission electron
microscopic levels.
30
The albino ferret lacks both retinal and
choroidal pigmentation.
30
The ferret has a tapetum cellulo-
sum situated in the superior choroid immediately external to
the choriocapillaris.
27
It contains rodlets that have a diame-
ter of 1.3–1.5 µm and a length of 3–4 µm (Table 2).
27
The
ferret tapetum lucidum is 5–7 cell layers arranged as a brick-
like structure (Table 2). Over the tapetum, the retinal
epithelium is nonpigmented.
30
Several layers of pigmented
melanocytes are present outside the tapetum lucidum in the
choroid of pigmented ferret eyes. A high concentration of
zinc cystein is found in the tapetal rodlets of ferrets.
Ranch mink (Mustela vison) All carnivores are believed to
possess a well-developed choroidal tapetum cellulosum.
3,27
However, the tapetum present in the superior part of the
fundus of the ranch mink seems to be degenerative. The
presence of only 2–3 layers of apparently degenerative tap-
etal cells containing a haphazard arrangement of reflective
material makes it very unlikely to function as an effective
tapetum lucidum (Table 2). The few tapetal cells demon-
strate a nonregular internal arrangement of membranes and
apparently degenerative structures that differs greatly from
the very regular, repeating pattern of reflective rodlets nor-
mally present within the cells of a developed tapetum cellu-
losum (Fig. 6b). The cause of this tapetal degeneration is
© 2004 American College of Veterinary Ophthalmologists, Veterinary Ophthalmology, 7, 11–22
+a r r +t· r tci rt· a ·oxc v r r +r r r a +r s 19
unknown but could be related to dietary insufficiencies in
specific animals, or perhaps the genetics of inbreeding.
3
Mammals – rodents
Paca (Cuniculus paca) A tapetum has only been reported in
one rodent; the paca or spotted cavy (Table 2). The tapetum
lucidum of the paca consists of several layers of ovoid-to-
flattened cells within the superior choroid.
32,33
The electron-
dense tapetal rodlets are typically uniform in diameter and
highly organized in carnivores, but vary in size, shape, and
orientation in the paca. Because the reflective material appears
to be not highly ordered regarding size, orientation and
spacing between the reflective units, the tapetum lucidum of
the paca is unlikely to provide constructive interference and
therefore produce a diffuse reflectance.
13
Rat (Long Evans rat) A vestigial tapetum has been sug-
gested in the hooded rat (Long Evans rat) for two reasons:
this tapetum is localized in the superior hemisphere and the
melanosone counts are very low in this dorsal area (Fig. 2,
lane 1, #3).
34
Mammals – cetacea
Grey seal (Halichoerus grypus) In the gray seal the tapetum
lucidum is located in the choroid and overlays the entire
effective fundus. It is composed of 30–35 layers of flattened
polygonal cells at its thickest and is reduced to 15–20 layers
of cells at the extreme periphery of the retina (Table 2). The
thick tapetum and the fact that it overlays the entire fundus
may be the result of the more three-dimensional environ-
ment in where this aquatic mammal resides. The tapetal cells
show less reflective material within their cytoplasm at the
scleral border of the tapetum, and are gradually replaced by
melanin-containing choroidal cells. The bulk of the cytoplasm
of the tapetal cells is filled with numerous membrane-bound
rodlets of reflective material of unknown composition. These
rodlets are 5–6 µm in length and 0.1 µm in width. They are
arranged in an organized hexagonal-lattice pattern with a space
of about 0.15 µm separating adjacent rodlets (Table 2).
4
Tapetum lucidum fibrosum
All hoofed animals or ungulates, as well as whales (Fig. 8d),
some marsupials, and one monkey (Aotes) are reported to possess
a tapetum fibrosum.
13,35,36
In ungulates, closely and regularly
arranged collagen fibers comprise the tapetum, which is
often referred to as a fibrous tapetum (Fig. 1). The fibrous
tapetum is basically acellular except for occasional fibrocytes
(Fig. 2). The collagen fibrils are organized into well-ordered
lamellae that branch and interconnect with adjacent lamellae
at the same level, being parallel with the retinal surface.
2
Cows The bovine tapetum fibrosum is situated in the
choroid and restricted mainly to the superior fundus.
36
Over
the central and mid-region of the tapetum the retinal
epithelial layer is completely unpigmented, whereas over
the peripheral tapetum, and certainly in the nontapetal
periphery of the retina, the RE is moderately to heavily pig-
mented. The bovine tapetum appears blue in color (Fig. 2,
lane 4, #4). The actual reflective material of the bovine tape-
tum is a large array of extracellular collagen fibrils arranged
in lamellae of varying thickness. Fibrocytes are located at
the borders of these lamellea. To date, collagen is the only
fiber type reported in tapeta fibrosa. The collagen fibrils are
arranged parallel to the surface of the tapetum, at a right
angle to incoming light. The collagen fibrils display the typi-
cal cross-striations of native collagen at about 640 Å, and
are fairly uniform in diameter at about 0.2 µm. They seem to
be arranged in a flattened hexagonal array, with individual
fibrils being separated by a center-to-center spacing of about
0.2 µm (Table 2). At the retinal edge of the tapetum, a rich
vascularized lamella containing the choriocapillaris separates
the tapetum from Bruch’s membrane. Blood vessels from the
pigmented outer region of the choroid penetrate the tape-
tum fibrosum to supply the choriocapillaris adjacent to
Bruch’s membrane. These penetrating blood vessels pass
through the tapetal region parallel to the incoming light
very close to a right angle. The bovine choriocapillaris,
while abutting directly against Bruch’s membrane, does not
indent this membrane nor the retinal epithelial layer.
36
Sheep and goat The sheep’s tapetum reflects a greenish blue
color in contrast to the adjoining regions of dense black pig-
mented choroid.
37
The tapetum is essentially a horizontal
strip in shape, and its lower edge just touches the point of
entry of the optic nerve (Fig. 2, lane 4, #2). The upper edge
is a more complex shape, and is less clearly defined. The
tapetum consists of a layer of collagen fibrils (100–200 µm
in depth), in which a few fibroblats are present (Fig. 8b).
23
These collagen fibers are 150 nm in diameter and are regu-
larly arranged with cross banding, in layers. Several hundred
of the lamellae or layers (600–740 layers) are estimated to
comprise the fibrous tapetum of the sheep.
37
Individual
layers of fibrils all run in a longitudinal direction, and are
arranged in a hexagonal pattern with the direction of light at
right angles to a side of the hexagon. The layers of fibrils are
frequently separated by cells which lie parallel to the retinal
surface (pigment cells, fibroblasts, smooth cells). There is a
maximum of 140 rows of collagen separated by 9–16 cellular
layers. The fibrils within a row are separated from each other
by a space of 67 nm, and the rows are separated from each
other by a space of 80 nm
2,23,37
The spacing between the
fibrils is 180 nm and does not seem to change with depth
(Table 2). The spacing between the rows appears to be more
constant than the diameter of the fibrils. In the nontapetal
areas, melanosomes are numerous, whereas over the central
tapetum fibrosum they are absent.
38
The goat has a tapetum similar in shape to the sheep
(Fig. 2, lane 4, #3). The penetrating blood vessels are less
uniformly arranged than in the cow, and even less than those
of the cat and dog tapetum.
23
20 or r i v i r r E T A L .
© 2004 American College of Veterinary Ophthalmologists, Veterinary Ophthalmology, 7, 11–22
Horses The fibrous tapetum in the horse is composed of a
layer of bundles of laminated areolar (loose) connective tis-
sue with an iridescent surface. There are four or five layers of
these fibers. The central tapetum of the horse is up to 30
microns thick (Fig. 8a). It extends horizontally from the sur-
face of the optic disc almost to the equator, and to a compa-
rable distance vertically above the optic disc to assume a
rounded triangle or semicircle (Fig. 2, lane 4, #1 and 2). The
choroidal capillaries penetrate the tapetum at right angles to
the collagen lamellae and they are called ‘Stars of Winslow’
when observed end-on ophthalmoscopically.
39
DISCUSSION
Tapetal color varies with species, breed, age, coat color and
amount of pigmentation in the eye and skin (Fig. 2). Structure,
organization and composition of the tapetum also vary from
species to species (Tables 1 and 2). Despite these differences,
however, all act to increase retinal sensitivity by reflecting
light back through the photoreceptor layer.
2
Physical properties of the tapetum lucidum: reflectivity
The high light reflectance of a tapetum is the result of thin-
film interference. The tapetal structure consists of several
alternative layers of high and low refractive index material
whose thicknesses are compatible with the wavelength of
light (one quarter of the wavelength of the light is the ideal
thickness and it would give the best reflectivity). At each
interface in such a structure, a certain proportion of the inci-
dent light is reflected and, depending on the thickness of the
layers and the wavelength being considered, the contribu-
tions from each interface interfere either constructively or
destructively.
19
The problem of producing efficient reflec-
tion in a biologic context is most often solved by arranging
material with a higher refractive index in a medium with a
lower refractive index. If the refractive material is randomly
arranged, diffuse reflectance will occur. If, however, the
higher refractive index material is of uniform size, orderly
arranged, and separated by specific and precise spacing, the
small number of reflectances will sum in a more constructive
manner and a higher overall reflectance will be achieved.
36
Denton has indicated that in an ideal theoretical model of
constructive interference, five layers of reflective material
would give close to 75% reflection.
14
Land states that for
most biologic systems the reflectance will approach 100%
after 10–20 layers.
19
The difference between actual and ideal
reflectance would seem to indicate that constructive inter-
ference in a biologic context is far from perfect. Structures
such as bloods vessels, nuclei of cells, and imperfections in
spacing and thickness of the reflective material probably all
interfere with reflectance, and therefore more layers are pre-
sumably required to create an effective tapetum lucidum.
5,17
From these different studies it appears that the extracellular
structures in the tapetum fibrosum, and the endocellular
ones in the tapetum cellulosum and retinal tapetum are iden-
tical in their spatial arrangement (Fig. 1). It therefore seems
that tapetal types have the same general reflective mecha-
nism regardless of their composition (Tables 1 and 2). It is
interesting to note that the cellular rodlets present substan-
tial chemical differences, compared to the collagen fibrils.
The diverse chemical nature of the different tapeta may be
responsible for determining different refractive indices. This
factor may play a role in the utilization of different wavelengths
of light by the animals according to their feeding behavior.
The hexagonal packing noted in the tapetum fibrosum and
in the tapetum cellulosum seems to be the most efficient
method to closely assemble cylindrical structures like colla-
gen fibrils or rodlets of reflective material in order to achieve
the best spatial arrangement. The relatively small differences
recorded in diameter and spacing may indicate a small differ-
ence in the optimal wavelength of reflection between theses
species, or might be due to the specimen fixation.
Specifically, the observed dimensions and spacing of the
collagen fibrils found in the bovine tapetum are commensu-
rate with the principles of constructive interference. The
dimensions and spacing observed for the collagen fibrils in
the bovine eye are similar to those noted in the sheep and
goat. The dimensions and spacing of collagen fibrils in the
tapetum fibrosum are largely similar to the dimensions and
spacing of the reflective rodlets in the tapetum cellulosum of
the cat, ferret and dog.
3,36
Nevertheless, spatial variations exist. In the cat, the tapetal
cells contain more precisely packed rodlets than those in the
ferret and dog (Fig. 7). The fact that the cat has a relatively
high number of layers (15–20) coupled with a great efficiency
of packing may explain the very great reflectance of the
feline tapetum cellulosum.
5
Rodieck has indicated that the
cat’s eye reflects about 130 times more light than the human
eye.
40
The dog tapetum, besides reflecting light of a different
wavelength, does so less efficiently. While the reflecting rodlets
of the dog are similar in dimensions to those of the cat
(Table 2), they appear to be more closely packed, resulting in
a unique value for the wavelength of maximal reflectance. In
addition, the canine tapetal rodlets are less precisely oriented
than those of the cat, which may result in a less efficient
reflectance from the dog tapetum (Fig. 7). How much more
light the feline tapetum reflects than the canine tapetum is
not known.
The thickness of the tapetum cellulosum varies also from
species to species, as described earlier (Table 2). The rela-
tively small differences in rodlet diameter and spacing may
indicate a small difference in the optimal wavelengths of
reflection between these species. While species differences
certainly exist with regard to tapetal thickness and rodlet
composition and organization, the intraspecific differences
that have been observed are probably due to breed variance,
sampling of different areas and indeed to developmental
stages.
In all species with a choroidal tapetum (either cellulosum or
fibrosum), the retinal epithelium overlying at least the cen-
tral region of the tapetum is nonpigmented (Fig. 1). This
occurrence obviously allows for free passage of light to
© 2004 American College of Veterinary Ophthalmologists, Veterinary Ophthalmology, 7, 11–22
+a r r +t· r tci rt· a ·oxc v r r +r r r a +r s 21
and from the tapetum. Pigment in this location would be
counterproductive to the function of a tapetum.
Among the vertebrates reported with a retinal tapetum are
a number of teleosts, crocodilians, the fruit bat, and the
opossum (Fig. 1). For these species, the tapetal material is
located within the retinal epithelium itself, hence positioning
the reflecting material closer to the photoreceptor cells.
Origin of the tapetal cells in tapeta cellulosa
The origin of the tapetal cells is still unknown. Some authors
believe the cells come from differentiation of connective tissue
cells
41,42
and some think they are modified melanocytes.
29,43
Some
investigators have proposed that tapetal cells and choroidal
melanocytes have a common origin from the cells of the meso-
derm which invade the primitive optic cup. One piece of evidence
to support this theory is that tapetal cells adjacent to the
choroid frequently contain both rods characteristic of the
tapetum and the pigment granules common to the choroidal
melanocytes (Fig. 9). There is therefore coexistence of the
tapetal rodlets and melanin granules in the same cells.
6,8,43
Cytochemical data have shown that the rodlets observed
in the cat are composed of melanin, and their development
follows a similar course to that described in other melano-
genic systems. The rodlet formation involves similar enzy-
matic mechanisms to those concerned in the construction
of melanosomes.
29
There are striking morphologic resem-
blances between the structural features of the specific inclu-
sions in the tapetal cells and those of developing melanosomes
in melanocytes. However, the tapetal cells exhibit a new mel-
anogenic system in which the melaninic structures have a
constant diameter and a regular pattern. Tapetal cells repre-
sent uniquely differentiated and highly specialized choroidal
melanocytes. It is interesting to note that, by the combina-
tion of structural and compositional modifications of the
melanosomes and their arrangement in the tapetal cells,
these highly specialized melanocytes have adopted the
capacity to reflect light instead of absorbing it.
6,43
Chemical properties of the tapetum lucidum
The reflective material varies widely in chemical composi-
tion from species to species. The collagen is the reflective
compound in tapeta fibrosa. It has been evaluated that colla-
gen constituted 65% of the dry weight of the sheep tapetum,
with a significant amount of amino acids such as proline,
hydroxyproline, lysine and hydroxylysine.
37
Among the
reflective materials noted in tapeta cellulosa are guanine,
cholesterol, zinc cysteine, riboflavin, pteridine, and a variety
of lipids (Table 2). A high level of zinc (i.e. zinc cysteine) is
present in the tapetal cells of both the dog and ferret tapeta,
whereas the tapetal cells contain abundant riboflavin in the
cat and the lemur
26,28
and abundant sulfur in the paca.
26
Riboflavin absorbs light at a low wavelength and fluoresces
at 520 nm, which is approximately the maximum absorption
of the rhodospin (500 nm). Thus, the cat and lemur tapeta
appear to not only reflect light but amplify it into a more
useful range by fluorescence.
5
The diverse chemical nature of the different tapeta may be
responsible for determining different refractive indices. This
factor may play a role in the utilization of different wave-
lengths of light by carnivores and herbivores. In fact, it is
well known that carnivores spend most of their active life at
night (crespucular), while herbivores lead a more diurnal life
(arrhythmic).
1,17
Vascularization of the tapetum lucidum and related structures
The choriocapillaris consists of a single layer of large-caliber
capillaries which are heavily fenestrated. The choriocapilla-
ris noted in various species with a tapetum cellulosum is
deeply indented into the RPE layer over the tapetal area. As
the choriocapillaris is not indented into the RPE of species
with a tapetum fibrosum, it is thought that this indenting of
the choriocapillaris into the RPE over a tapetum cellulosum
is mainly due to the rigidity of the reflective material within
the tapetal cells.
3,5,25
This may indicate that the choriocapil-
laris is also involved in the metabolic support of the tapetum
which is relatively poorly vascularized.
4,36
The radial align-
ment of the tapetal penetrating capillaries appears to be the
best way to connect the choriocapillaris to both its supplying
and draining systems, because this results in the least inter-
ference with tapetal reflectivity.
13
The ‘Stars of Winslow’
specifically describe the choroidal capillaries that penetrate
the tapetum at right angles to the collagen lamellae and that
can be observed ophthalmoscopically in horses. There are
also choroidal capillaries that penetrate the tapetum at right
angles to the long axes of the tapetal cells in carnivores.
They can occasionally been ophthalmoscopically observed
in cats, but rarely in dogs because of the brightness of the
fundus and the lack of magnification with conventional
ophthalmoscopes.
In most vertebrates Bruch’s membrane is reported as a
pentalaminate structure: the basal lamina of the choriocapil-
laris, the basal lamina of the retinal epithelium, and an inner
and outer collagen layer with a central discontinuous elastic
layer. In some species with a choroidal tapetum, Bruch’s
membrane may be reduced to only the two basal laminae. In
teleosts Bruch’s membrane is normally trilaminate (basal
lamina of the choroid, basal lamina of the epithelium and a
thick central collagenous layer).
27
CONCLUSION
The tapetum lucidum represents a remarkable example of
neural cell and tissue specialization as an adaptation to a dim
light environment. This adaptation appears to have evolved
on a number of separate occasions, and it is therefore not too
surprising that both its location and structure, as well as the
choice of reflective material are varied.
13
Among mammals, variations are greatest within the carni-
vore family. These variations, which are entirely cellular in
nature, may represent selective visual adaptations associated
with their feeding behavior. These adaptations are probably
in response to the use of specific wavelengths and amount of
22 or r i v i r r E T A L .
© 2004 American College of Veterinary Ophthalmologists, Veterinary Ophthalmology, 7, 11–22
reflectance required. Herbivores, on the other hand, appear
to have evolved less remarkably with regard to the tapetum,
having variations primarily directed to the amount of
reflectance.
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rods in the tapetal cells of the cat. Journal of Anatomy 1978; 127:
505–513.
30. Tjalve H, Frank A. Tapetum lucidum in the pigmented and albino
ferret. Experimental Eye Research 1984; 38: 341–351.
31. Braekevelt CR. Fine structure of the retinal epithelium and tape-
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1989; 135: 296–302.
32. Braekevelt CR. Fine structure of the tapetum lucidum of the paca
Cuniculus paca. Acta Anatomica (Basel) 1993; 146: 244–250.
33. Silveira LC, Picanco-Diniz CW, Oswaldo-Cruz E. Distribution
and size of ganglion cells in the retinae of large Amazon rodents.
Visual Neuroscience 1989; 2: 221–235.
34. Howell WL, Rapp LM, Williams TP. Distribution of melano-
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35. Braekevelt CR. Retinal epithelial fine structure in the vervet mon-
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36. Braekevelt CR. Fine structure of the bovine tapetum fibrosum.
Anatomia, Histologia, Embryologia 1986; 15: 215–222.
37. Bellairs R, Harkness ML, Harkness RD. The structure of the
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38. Braekevelt CR. Fine structure of the choriocapillaris, Bruch’s mem-
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39. Prince JH. Anatomy and Histology of the Eye and Orbit in Domestic
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40. Rodieck RW. The vertebrate retina. Principles of Structure and
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41. Hebel R. Development and structure of the retina and the tapetum
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42. Vogel H. Postnatal development of the cat’s retina. Advances in
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reptiles) (Table 1). 1).16 R E TIN AL TAPETU M L U C I DU M Retinal tapeta lucida are common in fish and some reptiles. a choroidal gland. the tapetum of gars possesses a yellow reflective material (yellow pigment) that occurs in regular array and is contained in a myriad of small spheres.10. Despite these differences. The presence of a tapetum lucidum influences the light sensitivity capabilities of a species. like the toadfish (Toadfishes).12.11 In contrast.10–12 The stratum argenteum is a ‘silvery’ reflective layer in the outer choroid. The tapetum of the seatrout (Sciaenidae) contains highly refractive lipid (triglycerides) particles.2.19 Animals without a tapetum lucidum (e. thereby enhancing visual sensitivity (Fig. red kangaroo and pig) generally have diurnal habits. It normally functions at low light levels to provide the light-sensitive retinal cells with a second opportunity for photon-photoreceptor stimulation. squirrels. the tapetum lucidum may harm visual acuity in bright illumination. and one or two choroidal glands.4.13 It is thought that the function of the occlusible tapetum is related primarily to reduction of ‘eye-shine’ rather than modifying retinal sensitivity. histologic and biochemical aspects of the tapetum lucidum of various vertebrates. 7. Tapeta lucida can be classified according to the location as well as the nature of the reflective materials that constitute them. and characterizes the tapetum of these teleosts as occlusible (Fig. as the reflective material is not organized within spheres. The phenomenon of ‘eye-shine’ is seen in a variety of animal species. adjacent to the sclera. 1). highly refractive material (tapetal rodlets) (Fig.9 In most of these species the retinal tapetum can be classified as either a lipid (in some teleosts. Veterinary Ophthalmology.11. as the tapetal reflection contributes only 30% of the light reaching the photoreceptors (70% of the light directly reaches the photoreceptors). or a layer of cells (which compose the tapetum lucidum cellulosum) packed with organized. an array of extracellular fibers (tapetum lucidum fibrosum). 1). but rarely occur in mammals.5.12 The choroid of toad fish is thin and contains blood vessels. and is generally thought to be related to the presence of an intraocular reflecting structure. 3). lane 1). A choroidal tapetum of reflecting cells lying immediately outside the choriocapillaris is formed in a few primitive teleosts. and a red or orange to pale gray fundus reflection (Fig.4 Tapeta lucida in vertebrates exhibit diverse structure. The reflecting material may be located within the cytoplasm of the retinal epithelium (retinal tapetum lucidum) or in the choroid. but because of a visual cell movement called the retinomotor movement. Melanin migrates towards the vitreous when the eye is illuminated and towards the sclera (backwards) in dim light or darkness (Fig.12 The reflective material is contained in spheres closely packed together in the apical processes of the pigment epithelial cells. 2). 11 The ‘eyeshine’ varies in color (from yellow to dark brown) depending on the retinomotor movement on the stage of light adaptation of the eye.7.3 The tapetum lucidum is a biologic reflector system that is a common feature in the eyes of vertebrates.10–12 In certain species the tapetum is occlusible not because of a pigment migration within a cell. 2. There is loose © 2004 American College of Veterinary Ophthalmologists. It can vary in chemical nature (Table 1).14.10. organization and composition (Fig. The choroidal tapetum lucidum lies immediately adjacent to the choriocapillaris and external to the retinal epithelium (Fig.g. and reflects light diffusely. and the other that lies in the pigment epithelium of the retina such as occurs in teleosts. 1). the tapetum lucidum is absent or vestigial and there is little choroidal melanin.1. The reflective material of the choroidal tapetum lucidum may be a guanine tapetum.12 whereas the tapetum of gars (Lepisosteidae) is composed of a yellow phenolic compound. the retinal tapetum lucidum.6.12         ET AL.6 This paper is a review of the anatomic. frequently present in fish larvae but not as frequent in adults. 11–22 .3. birds. The size of the eye and those of its different structures influence the amount of light gathered. and a stratum argenteum.14 The choroid of teleosts is a connective tissue layer rich in blood vessels. or nonocclusible (or static) (found in mammals). as well as the migration of the processes of the RPE cells between the cells and over the reflective material (elasmobranchs).11 The lipid tapetum observed in the seatrout has a white appearance. primates. Another differentiation can be made among the retinal tapeta lucida based on whether the tapetum is occlusible (found in most of teleosts).3. A B SEN C E O F T APETU M in teleosts is caused by the migration of black pigment (melanin) within the retinal pigment epithelium (RPE) and the choroidal cells. the tapetum lucidum is an integral part of the pigment epithelium in primitive teleosts (lying in the processes of the pigmented epithelial cells) (Fig.11.5 While increasing retinal sensitivity in dim illumination. all tapetal variants act to increase retinal sensitivity by reflecting light back through the photoreceptor layer. The migration of melanin can mask or unmask the reflecting material depending upon the degree of environmental illumination. More commonly. and the choroidal tapetum lucidum. The variation in ‘eye-shine’ in occlusible tapeta Fish – teleosts Two morphologic types of tapeta lucida can be observed in the eyes of fishes: one that lies in the choroid such as occurs in elasmobranchs (that will be discussed later). mammals) or guanine type (in some teleosts. In species known to have a stratum argenteum as adults. 1).2 Birds are apparently the only large group of animals in which the tapetum is consistently absent. Three basic tapetal morphotypes of vertebrates are described and compared: the absence of tapetum. The spheres are organized in parallel sheets with packing being hexagonal in the plane of the sheet. the tapetum lucidum. of the retina and the optical system.8 The red to orange background results from reflection of light from choroidal blood vessels.

The thin crystals are superposed over each other and form a good reflective system. r & c. The retinal tapetum lucidum in fish (teleosts): oscar (Astronotus oscellatus). inner nuclear layer. inner nuclear layer. original magnification ×400. sclera. 11 – 22 . original magnification ×200. fruit bat. (H&E staining. rods and cones. deer. sc. From right to left: lane 1: absence of tapetum: bird.) Figure 2. The retinal tapetum lucidum in alligator (Alligator mississipiensis). pig. hooded rat. onl. r & c. outer nuclear layer. and becomes a layer of pentameric crystals and a layer of yellow birefringent granules. onl. choroid. dog (3). Reflection on the tapetum lucidum. and is mostly a specular reflector. outer nuclear layer. ch. horse. tapetum /retinal pigment epithelium. t /rpe. It consists of two or more cell layers (iridophores). First stimulation on the rods and cones. Figure 5. human. Melanocytes are absent except for a few near the globe equator. The iridophores are polygonal in shape and loaded with packets of elongated thin pentagonal crystals that are oriented with their long axis parallel to the argenteal surface. (H&E staining. ch. is bilaminate. mouflon. Figure 4. inl. cow. original magnification ×200. sclera. The retinal tapetum lucidum of a fruit bat (Pteroptus giganteus). reflecting much of the incident light. t /rpe. Lane 2: retinal tapeta: alligator. cat.) connective tissue between the choriocapillaris and the stratum argenteum. leopard. sc. jaguar. inl. rods & cones. 3. tapetum /retinal pigment epithelium. albinos rat. 1. choroid. Incoming light ray. goat. red kangaroo. 4. r & c.) Figure 3.                              13 Figure 1. rhesus monkey. The outer surface of the argenteum is covered by a thin yellow sheet that has a mat appearance. Diagram of the three types of the tapetum lucidum present among vertebrates. sheep. t/rp. Lane 3: Choroidal tapeta cellulosa: puppy. The yellow packets are in the form of 1-µm diameter small © 2004 American College of Veterinary Ophthalmologists. Lane 4: choroidal tapeta fibrosa: foal. has an orange metallic luster. insert ×400. outer nuclear layer. Second stimulation on the rods and cones. ch. tapetum/retinal pigment epithelium. onl. possum. The inner surface of the stratum is shiny. choroid. 7. rods & cones. The reflective material is uric acid (Table 1). The outermost layer of the choroid is occupied by a thin stratum argenteum which lies adjacent to the sclera. Ocular fundus of various species showing the absence or the presence a tapetum lucidum that can exhibit various colors. Veterinary Ophthalmology. 2. (H&E staining.

9.) Figure 7. By moving posteriorly or anteriorly within the cells. Electronic microscopy of the tapetal area in a dog (a) and a cat (b). rpe. outer segment.9. both guanine crystals and melanosomes are found within the epithelial cells. inner nuclear layer. a semicircular area of the retinal epithelium in the superior fundus shows a fair sulfur yellow coloration and is specialized as a tapetum lucidum (Fig. Figure 6.15 © 2004 American College of Veterinary Ophthalmologists. t. The tapetum lucidum cellulosum consists of layers tapetal cells regularly arranged. onl. 4). The crystals measure 100–120 nm in thickness and up to 0. retinal pigment epithelium. appear to have a geometric form. inl. the tapetum is an occlusible type. no melanosomes are formed indicating a nonocclusible tapetum in the Caiman sclerops. 11–22 . the fine structure of the retinal pigment epithelial cells of the crocodilians in general resembles that found in other vertebrates (Fig. os.12 Reptiles – crocodilians In crocodilians. c.15. ch. the tapetum is occlusible in toadfish. and are packed closely in a continuous mass. Outer segments of the retinal receptors are deeply inserted into broad. capillary. choroid. no such regular orientation of the guanine has been seen in the spectacled caiman (Caiman sclerops). (Trichrome staining. tc. t. #1).15 Guanine crystals or calcium salts of guanine appear to be the most common reflective material in this tapetum lucidum (Table 1). and the cougar (d).9. sc. tapetal cell. rods & cones. Veterinary Ophthalmology. c. 1) and retina.14         ET AL. 7.15 At the edge of the tapetum. however. Tapetum cellulosum of the dog (a). retinal pigment epithelium. tapetum. sclera. Therefore. insert ×400. tapetum. mainly crystalcontaining processes of the pigment epithelial cells. The rodlets in the tapetal cells are more precisely oriented in the tapetum of the cat than in the tapetum of the dog.9 In contrast. lane 2. (Original magnification ×4000. which may allow dispersion of the reflected light to occur more readily. rpe. capillary.15 In the Caiman crocodilus the guanine crystals are highly organized at a cellular level.9 Other than the presence of guanine crystals. being perpendicular to the long axis of the retinal photoreceptors (e. the mink (b).15 In comparison. r & c.) granules. original magnification ×200. the reflecting material is much closer to the photoreceptors. therefore.15 In spectacled caiman.16 Nearly all of the epithelial cells contain membrane-bound tapetal crystals. to the path of light). In the tapetal area. outer nuclear layer. both the choroidal and the retinal pigment prevent the light from reaching the argenteum (Fig. 2.7 µm in length.g. the dark coloration of the inferior part of the fundus in the Caiman crocodilus is caused by masking of tapetal crystals by melanosomes. the cat (c).

11. inl. and the other which occurs in teleosts and lies in the pigment epithelium of the retina. very few or no melanosomes are found. The reflecting material consists of a Guanine tapetum Fish – elasmobranchs Two morphologic types of tapetum lucidum can be observed in the eyes of fish: one in the choroid that occurs in elasmobranchs. t. ch. #2).17 The reflecting material contains cholesterol or cholesterol esters (Table 1). indicating a nonocclusible tapetum. sclera. which corresponds to the measured reflectance peaks around 470 –520 nm (Table 2).18 C HO R O I D A L T A P E T U M LU C ID U M Figure 9. outer nuclear layer. This type of biologic photon reflector system approximates ‘ideal’ multilayer reflectors. 2. the epithelial cells contain both reflecting material and melanosomes. The tapetal material lies at the base of the retinal cells (which are arranged in a zig-zag manner) on the inner side of the cones of pigment (Fig. inner nuclear layer. The reflecting material is believed to be contained in many spherules and to be phospholipid in composition (Table 1). 5). 7. melanin granules.11.17 Mammals – fruit bats The tapetum lucidum in fruit bats (Pteroptus giganteus) appears white to yellow when isolated in vitro (Fig. The reflectance © 2004 American College of Veterinary Ophthalmologists. but the reflected color is red in vivo. c. insert ×400. (Trichrome staining. retinal pigment epithelium. original magnification ×200.1. r & c. tapetum.16. lane 2.14 The elasmobranch tapetum in Squalus and Scyliorhinus consists of sheets of 12–13 superposed guanine crystals in the choroid. rpe. the sheep (b). lane 2. The size and shape of the lipoidal spheres vary somewhat from cell to cell. The choriocapillaris over the tapetal area is organized in a similar way to other mammals. At the periphery. A tapetal cell in the tapetum cellulosum of the mink containing melanin granules (m). c. probably due to retinal blood.16. and the pilot whale (d). (Trichrome staining.16. rods & cones. Melanocytes may be the progenitor cell of the tapetal cells. onl. although three features weaken its capability. tapetal cell. capillary. 2. original magnification ×1000. rpe. Tapetum fibrosum of the horse (a).17 Centrally in the tapetal area. retinal pigment epithelium.13. sc.1.) Mammals – marsupials – opossum A tapetum lucidum is present in a semicircular area of the superior retinal epithelium of the opossum (Didelphis virginiana) (Fig. Veterinary Ophthalmology. #3). The tapetum lucidum fibrosum consists of layers of collagen fibers regularly arranged. as will be discussed later.1.                              15 Figure 8. choroid. The thickness of the crystals ranges from 60 to 80 nm. capillary. m. 11 – 22 . but all of the spheres within the same epithelial cell are generally of a similar morphologic appearance. the dolphin (c). tc.) large quantity of lipoidal spheres scattered throughout the epithelial cells.

and is oriented at an angle of 30° to incoming light. The pigment in retinal epithelial cells overlying the tapetum of the Squalus can migrate between and over the reflective cells during illumination. size) Occlusible Occlusible Argenteum stratum Occlusible Spherules triglycerides.21 These tapetal cells alternate with. and the presence of migrating retinal epithelial cells in Squalus (an occlusible tapetum) results in optical interference.1 µm in width and 5.4 µm Spherules yellow pigment (Phenolic compound). as light in the sea is very directional. anastomosing. and are separated from one another by melanocytes that extend beyond the tapetal cells to intervene between the tapetal cells and the incoming light. the tapetum cellulosum is composed of layers of rectangularshaped cells (Fig. known as the choriocapillaris. 1). diameter: 0.8 Over the tapetal region. pass through the tapetum lucidum at right angles and terminate in the extensive.3.8. The choroidal stroma adjacent to the tapetum lucidum contains numerous small blood vessels that penetrate the tapetal layer to form a single-layered capillary network. and retreat during darkness to expose the reflectors (Fig. A typical tapetal cell will have 15–20 layers of these reflective crystals throughout the thickness of the cell. being multilayered at its center. on the surface of the tapetum.20. shape. Retinal epithelial pigment migration may also occur to limit exposure of an animal with ‘eye-shine’.0–8. the arrangement of the reflective surface is not always parallel to the retinal plane. 1). The principal difference is in the high degree of order of the rods in the tapetal cells compared to the random packing of the melanin granules in the melanocytes. Table 1. At the periphery of the tapetum melanocytes often intermingle with tapetal cells. the retinal epithelium (RE) is nonpigmented to allow light to pass to and from the reflective tapetum. are heavily fenestrated and are separated from the tapetum or pigment cells of the choroid by a narrow © 2004 American College of Veterinary Ophthalmologists. These capillaries form a single layer which underlie all parts of the retina. cholesterol and esters˙ Spherules peak is much too broad. capillary network: the choriocapillaris.2 Histologically. The bulk of the tapetal cells are packed with evenly spaced guanine crystals. Tapetum lucidum cellulosum Mammals – carnivores Common features Many mammalian carnivores have a choroidal tapetum of a cellular type (Table 2).24 µm Acid uric Occlusible Non-occlusible Non-occlusible Spherules guanine Spherules guanine. the tapetal melanocytes are as thoroughly dominated by a single type of cytoplasmic component (melanin granules) as the tapetal cells. The elasmobranchs that have a choroidal tapeta often have very elaborate occlusible tapeta lucida. 7. 11–22 .22). diameter: 0. The crystals measure approximately 0.7 µm. and it is not uncommon to observe cells which display morphologic characteristics of both tapetal cells and melanocytes.16         ET AL.11 Gars11 Toadfish12 Reptiles Crocodiles Alligator mississipiensis13 Caiman crocodilus9 Caiman sclerops15 Marsupials Opossum16. At the periphery of the tapetum the RE becomes pigmented. The size and spacing of these reflective crystals is commensurate with the principles of constructive interference. Histologically.12 µm Guanine Lipoidal spherules. In nontapetal locations the RE is heavily pigmented to become the retinal pigment epithelium (RPE). and ‘eye-shine’ might increase the chance of being caught by predators. The thickness of the tapetum varies. and are separated by a space of 0.5.17 Placentals Bats Fruit bat (Eidelon)13.25 The layers of melanocytes gradually increase toward the periphery of the tapetum. and thinning to a single cell (or lamella) at its periphery and adjacent to the optic nerve. The tapetum cellulosum is thus interposed between the large branching vessels in the choroid and the single layer of the choriocapillaris beneath the retinal epithelium.16 Type of tapetum Reflective material (nature.23 –26 The tapetal cells are gradually replaced by melanincontaining cells of the choroid at the outer edge of the tapetum (Fig. The tapetum is supplied by branches of blood vessels of the pigmented choroid. as in some teleosts. 1).0 µm in length.25 µm (Table 2).6. These penetrating vessels run from the outer vascular layer of the choroid. diameter: 0. Veterinary Ophthalmology. Retinal tapeta among the vertebrates Classification Fishes Teleosts Seatrout10. length: 0. The retinal tapetum of the Port Jackson shark is an occlusible type. The choriodal tapetum of the Port Jackson shark (Hetrodontus phillipi ) consists of a single layer of overlapping cells situated immediately external to the choriocapillaris. These values are very similar to those found in other elasmobranchs (short-tailed stingray and southern fiddler ray 20.

15 µm Diameter: 8 – 10 µm Length: 0.30 µm Non regular Diameter: 5 – 6 µm Length: 0.15 µm Spacing: 0.10 – 0.08 µm Diameter: 5 µm Length: 0.08 µm Diameter: 0.15 µm Spacing: 0. Choroidal tapeta among the vertebrates Classification Fishes Elasmobranchs.20 µm Diameter: 4 – 6 µm Length: 0.18 µm Cat5. 11 – 22 .25 Cellulosum Riboflavin 43 µm. 2 – 3 layers 30 – 35 layers Ungulates Cow36 Fibrosum Sheep38 Fibrosum Collagen 100 – 200 µm 100 – 200 µm23 140 rows per layer: 9 – 16 layers26 Lemurs Lemur (Galago crassicaudatus)16 Rodents Paca (Cuniculus paca)32 Riboflavin Fibrosum ? 12–15 Diameter: 8 – 10 µm Length: 0.20 µm Diameter: 0.15 µm Spacing: 0. Veterinary Ophthalmology.18.20 µm16 Diameter: 0.10 µm Spacing: 0.2 – 0.25 µm Spacing: 0.15 µm Spacing: 0.23 µm Spacing: 0.                              17 Table 2. Zinc 16 – 20 layers26 30 layers8 18 – 30 layers23 6 –10 layers40 Ferret 26 Cellulosum Zinc cysteine Zinc cysteine 35 layers6 7 – 10 layers 5 – 7 layers30 50 µm.20. 10 – 12 layers41 15 – 20 layers Riboflavin.27 Type of tapetum Occlusible Occlusible Reflective material Guanine crystals Guanine Thickness.14 µm Spacing: 0.14.40 µm Diameter: 3 – 4 µm Length: 0. 10 – 15 layers27 Ranch Mink31 Pinnipedes Seal (Grey Seal)4 Cellulosum Cellulosum ? 30 µm.21 Placentals Carnivores Dog8. 7.18 µm Spacing: 0.13 – 0. number of cell layers or crystals 60 – 80 µm Length: 5 – 8 µm Width: 0.1 µm Size of fibrils or Rodlet’s dimensions 12 – 13 superposed crystals 15 – 20 superposed crystals Cellulosum Zinc cysteine 9 – 15 layers Zinc cysteine 9 – 11 layers26 15 – 20 layers3 Diameter: 4 µm Length: 0.15 µm Primates Non human (Aotus)16 Fibrosum Collagen © 2004 American College of Veterinary Ophthalmologists.15 µm Spacing: 0. Port Jackson Shark13.19 Elasmobranchs. Squalus13.20 µm Diameter: 0.15 µm Diameter: 0.20 µm Spacing: 0.15 µm Diameter: 6 µm Length: 0.12 µm Spacing: 0.20 µm Diameter: 4 µm Length: 0.

The bulk of the cell is filled with numerous. set in even brick-like rows.6.27 The ferret tapetum lucidum is 5–7 cell layers arranged as a bricklike structure (Table 2). 2).15 µm separating adjacent rodlets (Table 2).24. layer of connective tissue that extends over the retinal surface.16 µm. Sagittally. #2 –4).26 Tapetal cells appear as polygonal plates arranged much like ‘patio bricks’ within each layer.12 µm in diameter. with the exception of individuals with partial or full albinism. Though the tapetum is undetectable grossly in the new-born. the tapetum present in the superior part of the fundus of the ranch mink seems to be degenerative.25. The tapetum in the center of the tapetal fundus of the dog consists of 18–20 layers of cells (Table 2). the choriocapillaris is indented into the retinal epithelial layer leaving a flat inner surface to the reflecting tapetum. with a mean thickness of 0. 2. 2.2 µm (Table 2).27 However. 2).1–0. while in peripheral regions of the nontapetal fundus the RE is not indented by the choriocapillaris. the canine tapetum resembles a multiple layered ‘brick wall’ (Fig.25 These rodlets are grouped in bundles.5.28 µm. lane 3.g.30 Several layers of pigmented melanocytes are present outside the tapetum lucidum in the choroid of pigmented ferret eyes. Each cell is a rounded polygon. Fig. #5).18         ET AL.25–27 The tapetum lucidum develops late in the eyes of some animals (e. membrane-bound rodlets of reflective riboflavin (Fig. The number of cell layers gradually thins towards the periphery where it ends. and show an orderly arrangement (Fig. 7a). lane 3.6 However. Within the bundle rodlets are primarily arranged into a compact regular hexagonal array. the retinal epithelium is nonpigmented. Tapetal development initially produces the immature feline tapetum that is pale blue in color (Fig.5. the feline tapetal cells sometimes contain melanin granules and. similarly. and are precisely arranged in a hexagonal lattice pattern with a space of 0. 6a). 11–22 . Its predominant adult color varies from yellow to green between individuals (Fig.27. 2.0 µm and a diameter of 0.30 The albino ferret lacks both retinal and choroidal pigmentation.3 –1.6. A high concentration of zinc cystein is found in the tapetal rodlets of ferrets. Bruch’s membrane over tapetal areas is extremely attenuated. dogs and cats). The presence of only 2–3 layers of apparently degenerative tapetal cells containing a haphazard arrangement of reflective material makes it very unlikely to function as an effective tapetum lucidum (Table 2).5 µm and a length of 3–4 µm (Table 2).3.24. 6b). #1). In the tapetal and peritapetal areas.25 Histologically. with each layer containing approximately 15–18 ranks of rodlets.3. repeating pattern of reflective rodlets normally present within the cells of a developed tapetum cellulosum (Fig. with its apex upwards and its base passing horizontally just below the optic disc (Fig. some tapetal cell-like melanocytes contain tapetal rodlets.23–25 Aging results in a loss of tapetal layers such that the adult cat tapetum has fewer layers than the kitten.8.to 4-week-old kittens. The few tapetal cells demonstrate a nonregular internal arrangement of membranes and apparently degenerative structures that differs greatly from the very regular. the feline tapetum consists of 15–20 layers of cells in its center. The degree of pigmentation of the animal apparently has no effect on development of the tapetum. In the dog.3.24 At the border region of the tapetum. Animals born with mature eyes have a well-developed tapetum although some tapetal maturation occurs after birth.26 In sagittal planes the individual feline tapetal cells appear rectangular.14 µm. Within each bundle the centers of the rodlets are arranged parallel to the retinal surface with remarkable regularity (Fig. having essentially completed development at this time. and appear to contain a hard and apparently homogeneous core (Table 2). It is present in both the pigmented and albino eye. and is shaped roughly like a rounded equilateral triangle. 40 –48 µm in diameter and an average thickness of 3. progressively thins and eventually disappears towards the periphery (Table 2.6.26 It covers an area of approximately 30% of the superior fundus. In fresh preparations the reflected color of the canine tapetum varies from yellow-green to green-blue with an irregular marginal area (Fig. and is usually completely developed by 4 months of age (Fig. lane 3.5.28 The rodlets are 4–6 µm in length and 0.8. Subsequent enlargement of the eye causes the relative extent of the tapetum to be reduced. Ranch mink (Mustela vison) All carnivores are believed to possess a well-developed choroidal tapetum cellulosum. Ferret (Mustela putorius) The tapetum lucidum in the ferret is an iridescent triangular area in the rear pole of the eye above the optic disc. Only a small part of the tapetal cell is occupied by an ovoid and centrally located nucleus. The cause of this tapetal degeneration is © 2004 American College of Veterinary Ophthalmologists.3. it becomes obvious in 3. The cytoplasm is tightly packed with separate bundles of membrane-bound rodlets that displace other organelles and/or inclusions into the perinuclear or peripheral region (Fig.3 Individual tapetal rodlets are fairly constant in their dimensions with a mean length of 4. electron-dense.30 The ferret has a tapetum cellulosum situated in the superior choroid immediately external to the choriocapillaris.30 A comparison of pigmented and albino ferrets revealed no morphologic differences of their cellular tapeta at both light microscopic and transmission electron microscopic levels.5. 6c). appearing as a single homogeneous layer (although it is bilaminate). 6c). 7b).27 It contains rodlets that have a diameter of 1. Over the tapetum.29 Dog (Canis familiaris) The tapetum lucidum in the dog is a rounded equilateral triangle in shape. with a mean spacing of 0. feline tapetal cells (located close to the retinal epithelial layer) are flattened (thickness of 3–4 µm). having the largest surface parallel to the retina. 7b). which normally extend through the entire thickness of the cell.25. 7. with the base oriented horizontally (excluding the optic disc in most of the breeds) and the apex directed upwards. Cat (Felis catus) The tapetum lucidum in the cat covers nearly 50% of the fundus area. Veterinary Ophthalmology.

8b).38 The goat has a tapetum similar in shape to the sheep (Fig. The bovine choriocapillaris.2 µm (Table 2). 11 – 22 . being parallel with the retinal surface. whereas over the peripheral tapetum. It is composed of 30–35 layers of flattened polygonal cells at its thickest and is reduced to 15–20 layers of cells at the extreme periphery of the retina (Table 2). Several hundred of the lamellae or layers (600–740 layers) are estimated to comprise the fibrous tapetum of the sheep. which is often referred to as a fibrous tapetum (Fig.13. smooth cells). The actual reflective material of the bovine tapetum is a large array of extracellular collagen fibrils arranged in lamellae of varying thickness. the tapetum lucidum of the paca is unlikely to provide constructive interference and therefore produce a diffuse reflectance. These penetrating blood vessels pass through the tapetal region parallel to the incoming light very close to a right angle. the paca or spotted cavy (Table 2). fibroblasts. At the retinal edge of the tapetum.32. 2. some marsupials. The layers of fibrils are frequently separated by cells which lie parallel to the retinal surface (pigment cells. The thick tapetum and the fact that it overlays the entire fundus may be the result of the more three-dimensional environment in where this aquatic mammal resides. orientation and spacing between the reflective units. in layers.                              19 unknown but could be related to dietary insufficiencies in specific animals. and its lower edge just touches the point of entry of the optic nerve (Fig. in which a few fibroblats are present (Fig. shape. The collagen fibrils are arranged parallel to the surface of the tapetum. There is a maximum of 140 rows of collagen separated by 9–16 cellular layers. #3). These rodlets are 5 –6 µm in length and 0. but vary in size.13 Rat (Long Evans rat) A vestigial tapetum has been suggested in the hooded rat (Long Evans rat) for two reasons: this tapetum is localized in the superior hemisphere and the melanosone counts are very low in this dorsal area (Fig. The fibrils within a row are separated from each other by a space of 67 nm. while abutting directly against Bruch’s membrane.23.33 The electrondense tapetal rodlets are typically uniform in diameter and highly organized in carnivores. and are gradually replaced by melanin-containing choroidal cells. 2. 7. The spacing between the rows appears to be more constant than the diameter of the fibrils.37 The spacing between the fibrils is 180 nm and does not seem to change with depth (Table 2).15 µm separating adjacent rodlets (Table 2). and are fairly uniform in diameter at about 0. They seem to be arranged in a flattened hexagonal array.36 In ungulates. a rich vascularized lamella containing the choriocapillaris separates the tapetum from Bruch’s membrane. #4). 8d). To date. and the rows are separated from each other by a space of 80 nm2. at a right angle to incoming light.3 Mammals – rodents Paca (Cuniculus paca) A tapetum has only been reported in one rodent. and is less clearly defined. whereas over the central tapetum fibrosum they are absent.37 Individual layers of fibrils all run in a longitudinal direction. The bulk of the cytoplasm of the tapetal cells is filled with numerous membrane-bound rodlets of reflective material of unknown composition. In the nontapetal areas. Veterinary Ophthalmology. The collagen fibrils display the typical cross-striations of native collagen at about 640 Å. They are arranged in an organized hexagonal-lattice pattern with a space of about 0. #2). as well as whales (Fig.37 The tapetum is essentially a horizontal strip in shape. lane 4.2 Cows The bovine tapetum fibrosum is situated in the choroid and restricted mainly to the superior fundus. The tapetum lucidum of the paca consists of several layers of ovoid-toflattened cells within the superior choroid. melanosomes are numerous. The bovine tapetum appears blue in color (Fig. The tapetal cells show less reflective material within their cytoplasm at the scleral border of the tapetum. 2. and one monkey (Aotes) are reported to possess a tapetum fibrosum. The fibrous tapetum is basically acellular except for occasional fibrocytes (Fig. 1). and orientation in the paca. 2. The upper edge is a more complex shape. the RE is moderately to heavily pigmented. lane 4.36 Over the central and mid-region of the tapetum the retinal epithelial layer is completely unpigmented.2 µm. Because the reflective material appears to be not highly ordered regarding size. The penetrating blood vessels are less uniformly arranged than in the cow.35. closely and regularly arranged collagen fibers comprise the tapetum. #3). collagen is the only fiber type reported in tapeta fibrosa.23 © 2004 American College of Veterinary Ophthalmologists. or perhaps the genetics of inbreeding. Fibrocytes are located at the borders of these lamellea.4 Tapetum lucidum fibrosum All hoofed animals or ungulates. with individual fibrils being separated by a center-to-center spacing of about 0. Blood vessels from the pigmented outer region of the choroid penetrate the tapetum fibrosum to supply the choriocapillaris adjacent to Bruch’s membrane. and are arranged in a hexagonal pattern with the direction of light at right angles to a side of the hexagon.1 µm in width. and even less than those of the cat and dog tapetum. does not indent this membrane nor the retinal epithelial layer. 2).34 Mammals – cetacea Grey seal (Halichoerus grypus) In the gray seal the tapetum lucidum is located in the choroid and overlays the entire effective fundus. The collagen fibrils are organized into well-ordered lamellae that branch and interconnect with adjacent lamellae at the same level. lane 1.36 Sheep and goat The sheep’s tapetum reflects a greenish blue color in contrast to the adjoining regions of dense black pigmented choroid. lane 4.23 These collagen fibers are 150 nm in diameter and are regularly arranged with cross banding. and certainly in the nontapetal periphery of the retina. The tapetum consists of a layer of collagen fibrils (100–200 µm in depth).

14 Land states that for most biologic systems the reflectance will approach 100% after 10–20 layers.5. The thickness of the tapetum cellulosum varies also from species to species. The hexagonal packing noted in the tapetum fibrosum and in the tapetum cellulosum seems to be the most efficient method to closely assemble cylindrical structures like collagen fibrils or rodlets of reflective material in order to achieve the best spatial arrangement. resulting in a unique value for the wavelength of maximal reflectance. The dimensions and spacing observed for the collagen fibrils in the bovine eye are similar to those noted in the sheep and goat. however. In addition. however. 1). the retinal epithelium overlying at least the central region of the tapetum is nonpigmented (Fig. 7). besides reflecting light of a different wavelength. It therefore seems that tapetal types have the same general reflective mechanism regardless of their composition (Tables 1 and 2). coat color and amount of pigmentation in the eye and skin (Fig. It is interesting to note that the cellular rodlets present substantial chemical differences. compared to the collagen fibrils. which may result in a less efficient reflectance from the dog tapetum (Fig. depending on the thickness of the layers and the wavelength being considered. or might be due to the specimen fixation. The choroidal capillaries penetrate the tapetum at right angles to the collagen lamellae and they are called ‘Stars of Winslow’ when observed end-on ophthalmoscopically. diffuse reflectance will occur.2 Physical properties of the tapetum lucidum: reflectivity The high light reflectance of a tapetum is the result of thinfilm interference. breed. Structures such as bloods vessels.40 The dog tapetum.36 Denton has indicated that in an ideal theoretical model of constructive interference. spatial variations exist. The dimensions and spacing of collagen fibrils in the tapetum fibrosum are largely similar to the dimensions and spacing of the reflective rodlets in the tapetum cellulosum of the cat.36 Nevertheless.5 Rodieck has indicated that the cat’s eye reflects about 130 times more light than the human eye. This occurrence obviously allows for free passage of light to © 2004 American College of Veterinary Ophthalmologists. Horses The fibrous tapetum in the horse is composed of a layer of bundles of laminated areolar (loose) connective tissue with an iridescent surface. as described earlier (Table 2).3. and to a comparable distance vertically above the optic disc to assume a rounded triangle or semicircle (Fig. the observed dimensions and spacing of the collagen fibrils found in the bovine tapetum are commensurate with the principles of constructive interference. ferret and dog. does so less efficiently. the tapetal cells contain more precisely packed rodlets than those in the ferret and dog (Fig. It extends horizontally from the surface of the optic disc almost to the equator. the small number of reflectances will sum in a more constructive manner and a higher overall reflectance will be achieved. and the endocellular ones in the tapetum cellulosum and retinal tapetum are identical in their spatial arrangement (Fig. Veterinary Ophthalmology. If the refractive material is randomly arranged.39 DISC USSIO N Tapetal color varies with species. #1 and 2). a certain proportion of the incident light is reflected and.19 The difference between actual and ideal reflectance would seem to indicate that constructive interference in a biologic context is far from perfect. The relatively small differences recorded in diameter and spacing may indicate a small difference in the optimal wavelength of reflection between theses species. Specifically. 7). 7. This factor may play a role in the utilization of different wavelengths of light by the animals according to their feeding behavior. The tapetal structure consists of several alternative layers of high and low refractive index material whose thicknesses are compatible with the wavelength of light (one quarter of the wavelength of the light is the ideal thickness and it would give the best reflectivity). There are four or five layers of these fibers. they appear to be more closely packed. and therefore more layers are presumably required to create an effective tapetum lucidum. all act to increase retinal sensitivity by reflecting light back through the photoreceptor layer. How much more light the feline tapetum reflects than the canine tapetum is not known. and separated by specific and precise spacing. Structure. While species differences certainly exist with regard to tapetal thickness and rodlet composition and organization. Despite these differences. 8a). At each interface in such a structure. The diverse chemical nature of the different tapeta may be responsible for determining different refractive indices. If. The fact that the cat has a relatively high number of layers (15 –20) coupled with a great efficiency of packing may explain the very great reflectance of the feline tapetum cellulosum. lane 4. nuclei of cells. 2). and imperfections in spacing and thickness of the reflective material probably all interfere with reflectance. orderly arranged.20         ET AL. five layers of reflective material would give close to 75% reflection. organization and composition of the tapetum also vary from species to species (Tables 1 and 2). 1). the canine tapetal rodlets are less precisely oriented than those of the cat. the higher refractive index material is of uniform size. In the cat. the contributions from each interface interfere either constructively or destructively. The relatively small differences in rodlet diameter and spacing may indicate a small difference in the optimal wavelengths of reflection between these species. sampling of different areas and indeed to developmental stages. In all species with a choroidal tapetum (either cellulosum or fibrosum).19 The problem of producing efficient reflection in a biologic context is most often solved by arranging material with a higher refractive index in a medium with a lower refractive index.17 From these different studies it appears that the extracellular structures in the tapetum fibrosum. The central tapetum of the horse is up to 30 microns thick (Fig. 11–22 . While the reflecting rodlets of the dog are similar in dimensions to those of the cat (Table 2). age. the intraspecific differences that have been observed are probably due to breed variance. 2.

7. and the opossum (Fig. cholesterol. zinc cysteine) is present in the tapetal cells of both the dog and ferret tapeta.1. by the combination of structural and compositional modifications of the melanosomes and their arrangement in the tapetal cells.43 Chemical properties of the tapetum lucidum The reflective material varies widely in chemical composition from species to species. Thus.27 C O N C LU S IO N The tapetum lucidum represents a remarkable example of neural cell and tissue specialization as an adaptation to a dim light environment. Tapetal cells represent uniquely differentiated and highly specialized choroidal melanocytes.5. The choriocapillaris noted in various species with a tapetum cellulosum is deeply indented into the RPE layer over the tapetal area. pteridine. it is well known that carnivores spend most of their active life at night (crespucular). 9). the basal lamina of the retinal epithelium. It has been evaluated that collagen constituted 65% of the dry weight of the sheep tapetum.25 This may indicate that the choriocapillaris is also involved in the metabolic support of the tapetum which is relatively poorly vascularized. 11 – 22 . the tapetal material is located within the retinal epithelium itself. and an inner and outer collagen layer with a central discontinuous elastic layer.17 Origin of the tapetal cells in tapeta cellulosa The origin of the tapetal cells is still unknown. as well as the choice of reflective material are varied. but rarely in dogs because of the brightness of the fundus and the lack of magnification with conventional ophthalmoscopes. Bruch’s membrane may be reduced to only the two basal laminae. and it is therefore not too surprising that both its location and structure. Veterinary Ophthalmology. which is approximately the maximum absorption of the rhodospin (500 nm). hence positioning the reflecting material closer to the photoreceptor cells. There are also choroidal capillaries that penetrate the tapetum at right angles to the long axes of the tapetal cells in carnivores. One piece of evidence to support this theory is that tapetal cells adjacent to the choroid frequently contain both rods characteristic of the tapetum and the pigment granules common to the choroidal melanocytes (Fig. variations are greatest within the carnivore family. and a variety of lipids (Table 2).29. There is therefore coexistence of the tapetal rodlets and melanin granules in the same cells. The diverse chemical nature of the different tapeta may be responsible for determining different refractive indices. riboflavin. In fact. This adaptation appears to have evolved on a number of separate occasions. these highly specialized melanocytes have adopted the capacity to reflect light instead of absorbing it. with a significant amount of amino acids such as proline.13 The ‘Stars of Winslow’ specifically describe the choroidal capillaries that penetrate the tapetum at right angles to the collagen lamellae and that can be observed ophthalmoscopically in horses.37 Among the reflective materials noted in tapeta cellulosa are guanine. However. The rodlet formation involves similar enzymatic mechanisms to those concerned in the construction of melanosomes. Pigment in this location would be counterproductive to the function of a tapetum.13 Among mammals. crocodilians. lysine and hydroxylysine. For these species. because this results in the least interference with tapetal reflectivity. Some authors believe the cells come from differentiation of connective tissue cells41. In teleosts Bruch’s membrane is normally trilaminate (basal lamina of the choroid. may represent selective visual adaptations associated with their feeding behavior.5 Vascularization of the tapetum lucidum and related structures The choriocapillaris consists of a single layer of large-caliber capillaries which are heavily fenestrated. 1). In some species with a choroidal tapetum. and their development follows a similar course to that described in other melanogenic systems. while herbivores lead a more diurnal life (arrhythmic). the fruit bat.42 and some think they are modified melanocytes.6. basal lamina of the epithelium and a thick central collagenous layer).28 and abundant sulfur in the paca. the cat and lemur tapeta appear to not only reflect light but amplify it into a more useful range by fluorescence. These adaptations are probably in response to the use of specific wavelengths and amount of © 2004 American College of Veterinary Ophthalmologists. In most vertebrates Bruch’s membrane is reported as a pentalaminate structure: the basal lamina of the choriocapillaris.3.                              21 and from the tapetum. They can occasionally been ophthalmoscopically observed in cats. This factor may play a role in the utilization of different wavelengths of light by carnivores and herbivores. it is thought that this indenting of the choriocapillaris into the RPE over a tapetum cellulosum is mainly due to the rigidity of the reflective material within the tapetal cells.29 There are striking morphologic resemblances between the structural features of the specific inclusions in the tapetal cells and those of developing melanosomes in melanocytes.e.43 Some investigators have proposed that tapetal cells and choroidal melanocytes have a common origin from the cells of the mesoderm which invade the primitive optic cup. The collagen is the reflective compound in tapeta fibrosa. hydroxyproline.43 Cytochemical data have shown that the rodlets observed in the cat are composed of melanin.6. As the choriocapillaris is not indented into the RPE of species with a tapetum fibrosum. zinc cysteine.4. It is interesting to note that. which are entirely cellular in nature. the tapetal cells exhibit a new melanogenic system in which the melaninic structures have a constant diameter and a regular pattern.26 Riboflavin absorbs light at a low wavelength and fluoresces at 520 nm. These variations. whereas the tapetal cells contain abundant riboflavin in the cat and the lemur26.8.36 The radial alignment of the tapetal penetrating capillaries appears to be the best way to connect the choriocapillaris to both its supplying and draining systems. Among the vertebrates reported with a retinal tapetum are a number of teleosts. A high level of zinc (i.

Cranbrook Press. Some reflective properties of the tapetum lucidum of the cat’s eye. 6. Shek JW. 150: 213–217. Wen GY. Nicol JA. Development and structure of the retina and the tapetum lucidum in the dog. 14. Graefe’s Archive for Clinical and Experimental Ophthalmology 1990. 33. Journal of Anatomy 1983. A comparative study of the tapetum.) The eye in evolution. Anatomia. Fine structure of the tapetum cellulosum of the grey seal (Halichoerus grypus). Distribution and size of ganglion cells in the retinae of large Amazon rodents. 1942. Rodieck RW. 54: 114 – 121. Acta Anatomica (Basel) 1989. Baumgarten HG. Laboratory Animal Science 1985. 278 –619. The tapetal cell: a unique melanocyte in the tapetum lucidum cellulosum of the cat (Felis domestica L. 127: 505 – 513. 26. Braekevelt CR. Distribution of melanosomes across the retinal pigment epithelium of a hooded rat: implication for light damage. 3rd edn. Tapetum lucidum in the pigmented and albino ferret. The vertebrate retina. Vogel H. 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