CHAPTER 6: INTEGUMENT

The integument (or skin)

It is a composite organ. On the surface is the epidermis, below it is the dermis, and
between them lies the basement membrane (basal lamina and reticular lamina).

The integument is one of the largest organs of the body, making up some 15% of the
human body weight.

Specialized Functions of Integument

As the critical border between the organism and its exterior environment, the
integument has a variety of specialized functions.
- It forms part of the exoskeleton and thickens to resist mechanical injury.
- The barrier it establishes prevents the entrance of pathogens.
- Skin swabs from healthy human volunteers identified over 200 different genera
of resident bacteria, including many that, if given the chance through a breach in the skin, would
produce serious staph infections, acne, and eczema, among other pathologies.
- Helps hold the shape of an organism as well
- Osmotic regulation and movement of gases and ions to and from the circulation
are aided by the integument in conjunction with other systems.
- Skin gathers needed heat, or radiates the excess, and houses sensory receptors.
- It holds feathers for locomotion, hair for insulation, and horns for defense.
- Skin pigments block harmful sunlight and display bright colors during courtship

GENERAL FEATURES OF THE INTEGUMENT

The Composite Organ (What’s inside the Composite Organ?)

1. Epidermis (surface)
2. Dermis (below the surface)

I. Dermis
The dermis of many vertebrates produces plates of bone directly through
intramembranous ossification. Because of their embryonic source and initial position within the
dermis, these bones are called dermal bones. They are prominent in ostracoderm fishes but
appear secondarily even in derived groups, such as in some species of mammals.

* Fibrous connective tissue - most conspicuous component of the dermis composed of

mostly collagen fibers
*Collagen fibers - woven into distinct layers called plies.
 The dermis of the protochordate amphioxus
exhibits an especially ordered arrangement
of collagen within each ply (figure 6.4). In
turn, plies are laminated together in very
regular, but alternating, orientation. These
alternating layers act like warp and weft
threads of cloth fabric, giving some shape to
the skin and preventing it from sagging.

In aquatic vertebrates, such as sharks, the

bundles of collagen lie at angles to each
other, giving the skin a bias, like cloth; that
is, the skin stretches when it is pulled at an
angle oblique to the direction of the bundles.

In fishes and aquatic vertebrates (includes

cetaceans and aquatic squamates), collagen
fibers of the dermis are usually arranged in
orderly plies that form a recognizable
stratum compactum.

In terrestrial vertebrates, the stratum

compactum is less obvious because
locomotion on land depends more on the limbs and less on the trunk. And, of course, any
wrinkling of the skin is less disruptive to a terrestrial vertebrate moving through air.
Consequently, collagen fibers are present, even abundant, in the skin of terrestrial vertebrates,
but they are much less regularly ordered and usually do not form distinct plies.

II. Epidermis
The epidermis of many vertebrates produces mucus to moisten the surface of the skin.
Epidermis
Fishes (Produces) Mucus -protection from bacterial infection

-helps ensure the laminar flow of water across the

body surface
Amphibians (Produces) Mucus -serves similarly from fishes

-keeps the skin from drying during sojourn on land

Terrestrial (Forms) Stratum -covering the body
Vertebrates Corneum - an outer
keratinized or -help them address life in a drying and abrasive
cornified layer terrestrial environment (one of the tetrapods
innovations)
Sauropsids (Forms) Stratum -Alpha-keratins are present in most flexible
(reptiles and Corneum epidermal layers where shape changes occur.
birds)
 (Produces) -Beta-keratins are more common in specializations
Keratinocytes - such as hard scales, claws, beak, and feathers.
alpha (soft), beta
(hard) forms of
keratin
Synapsids (taxonomic subclass within Only alpha-keratins are present.
the class Reptilia)

The epidermis is derived from the ectoderm

and produces the basal lamina (figure 6.1a). The
dermis develops from mesoderm and
mesenchyme and produces the reticular lamina.
Between the integument and deep body
musculature is a transitional subcutaneous
region made up of very loose connective and
adipose tissues. In microscopic examination, this
region is termed the hypodermis. In gross
anatomical dissection, the hypodermis is referred
to as the superficial fascia (figure 6.1b).

Epidermis and dermis together

-Form some of the most varied structures found
within vertebrates: teeth, denticles, and scales of
fish.
-They are so closely linked across the basement
membrane.
PHYLOGENY
INTEGUMENT OF FISHES
-Fish integument is a large organ that is
continuous with the lining of all the body
openings, and also covers the fins. In
addition to its protective functions, fish skin
may serve important roles in
communication, sensory perception,
locomotion, respiration, ion regulation,
excretion, and thermal regulation.
-Two types of cells occur within the
epidermis of fishes: epidermal cells and
specialized unicellular glands. In living
fishes, including cyclostomes, prevalent
epidermal cells make up the stratified
epidermis. Superficial epidermal cells are
tightly connected through cell junctions and
contain numerous secretory vesicles that are
released to the surface where they
contribute to the mucous cuticle.
Epidermal cells of the basal layer are
cuboidal or columnar. Mitotic activity is
present in but not restricted to the basal
layer. Unicellular glands are single,
specialized, and interspersed among the
epidermal cell population. There are several
types of unicellular glands. The club cell is
an elongate, sometimes binucleate,
unicellular gland. Some chemicals within
club cells excite alarm or fear. They are
thought to be released by observant
individuals to warn others of imminent
danger. The granular cell is a diverse cell
found in the skin of lampreys and other
fishes. Both granular and club cells
contribute to the mucous cuticle, but their
other functions are not fully understood.
The goblet cell is a type of unicellular
gland that is absent from lamprey skin but
is usually found in other bony and
cartilaginous fishes.(6.7)
Primitive Fishes
-Dermal bones of the cranial region were
large, forming the head shields; but more
posteriorly along the body, the dermal
bones tended to be broken up into smaller
pieces, the dermal scales. The surface of
these scales was often ornamented with
tiny, mushroom shaped tubercles. These
tubercles consisted of a surface layer of
enamel or an enamel-like substance over an
inner layer of dentin. One or several
radiating pulp cavities resided within each
tubercle. The dermal bone supporting these
tubercles was lamellar, organized in a
layered pattern. The skin of living hagfishes
and lampreys departs considerably from
that of primitive fossil fishes. Dermal bone
is absent, and the skin surface is smooth
and without scales. The epidermis is
composed of stacked layers of numerous
living epidermal cells throughout. Within
the dermis, hagfishes also possess
multicellular slime glands that release their
products via ducts to the surface. (6.8)
Chondrichthyes
-In cartilaginous fishes, dermal bone is
absent, but surface denticles, termed
placoid scales, persist. These scales are what
give the rough feel to the surface of the skin
(figure 6.9a). Recent evidence suggests that
these tiny placoid scales favorably affect the
water flowing across the skin as the fish
swims forward to reduce friction drag.
Numerous secretory cells are present in the
epidermis as well as stratified epidermal
cells. The dermis is composed of fibrous
connective tissue, especially elastic and
collagen fibers, whose regular arrangement
forms a fabric like warp and weft in the
dermis. This gives the skin strength and
prevents wrinkling during swimming. (6.9)
Bony Fishes
-The dermis of bony fishes is subdivided
into a superficial layer of loose connective
tissue and a deeper layer of dense fibrous
connective tissue. Chromatophores are
found within the dermis. The most
important structural product of the dermis
is the scale. In bony fishes, dermal scales do
not actually pierce the epidermis, but they
are so close to the surface they give the
impression that the skin is hard. The
epidermal covering includes a basal layer of
cells. Above this layer are stratified
epidermal cells. As they move toward the
surface, epidermal cells undergo
cytoplasmic transformation, but they do not
become keratinized. Within these layered
epidermal cells occur single unicellular
glands, the secretory and club cells. These
unicellular glands, along with epidermal
cells, are the source of the mucous cuticle,
or surface “slime.” (6.10)
The ganoid scale is characterized by the
prevalence of a thick surface coat of enamel
(ganoin), without an underlying layer of
dentin (figure 6.11b). Dermal bone forms
the foundation of the ganoid scale,
appearing as a double layer of vascular and
lamellar bone (in palaeoniscoid fishes) or a
single layer of lamellar bone (in other
ancestral actinopterygians). Ganoid scales
are shiny (because of the enamel),
overlapping, and interlocking. Living
polypteriformes and gars retain ganoid
scales. However, in most other lines of bony
fishes, ganoid scales are reduced through
the loss of the vascular layer of bone and
loss of the enamel surface. This produces, in
teleosts, a rather distinctive scale. The
teleost scale lacks enamel, dentin, and a
vascular bone layer. Only lamellar bone
remains, which is acellular and mostly
noncalcified. Two kinds of teleost scales are
recognized. One is the cycloid scale,
composed of concentric rings, or circuli.
The other is the ctenoid scale, with a
fringe of projections along its posterior
margin. New circuli are laid down, like rings
in a tree, as a teleost fish grows. Annual
cycles are evident in the groupings of these
circuli, and from this pattern in the scales
we can determine the age of individual fish.
(6.11)
INTEGUMENT OF TETRAPODS
-Multicellular glands are more common in
the skin of tetrapods than in the skin of
fishes. In fishes, the mucous cuticle and
secretions of the unicellular glands at or
near the surface of the skin coat it. In
contrast, among tetrapods, multicellular
glands usually reside in the dermis and
reach the surface through common ducts
that pierce the cornified layer. Thus, the
stratum corneum that protects the skin and
prevents desiccation also controls the
release of secretions directly to the surface.
If it were not for these openings in the
stratum corneum, the surface of the skin
could not be coated or lubricated by these
secretions.
Amphibians
-Amphibians are of special interest because
during their lives they usually
metamorphose from an aquatic form to a
terrestrial form. In most modern
amphibians, the skin is also specialized as a
respiratory surface across which gas
exchange occurs, with the capillary beds in
the lower epidermis and deeper dermis. In
fact, some salamanders lack lungs and
depend entirely on cutaneous respiration
through the skin to meet their metabolic
needs. The most primitive tetrapods had
scales like the fishes from which they arose.
Among living amphibians, dermal scales are
present only as vestiges in some species of
tropical caecilians (Apoda). During the
breeding season, nuptial pads may form
on digits or limbs of male salamanders or
frogs. Nuptial pads are raised calluses of
cornified epidermis that help the male hold
the female during mating. The mucous
glands tend to be smaller, each being made
up of a little cluster of cells that release their
product into a common duct.
The poison glands (granular glands) tend
to be larger and often contain stored
secretions within the lumen of each gland.
Secretions of poison glands tend to be
distasteful or even toxic to predators.
However, few persons handling amphibians
are bothered by this secretion, nor need
they be concerned, because it is potentially
harmful only if eaten or injected into the
bloodstream. (6.12)
Reptiles
Reptiles show more advanced integumental
adaptations to the terrestrial environment
because they are more far-removed from the
water. In contrast, the cells are more highly
keratinized. The integument is modified
into horny scales in snakes and lizards. In
snakes, the scales on the ventral surface can
be further modified into scutes, which can
be used in locomotion. In turtles the
epidermis is strongly modified into plates
that cover the shell, and because they
increase in diameter each year, they can be
used to age the animals. Additionally,
epidermal scales may be modified into
crests, spines, or horn-like processes.
Although not usually associated with scales,
dermal bone is present in many reptiles.
The gastralia, a collection of bones in the
abdominal area, are examples. Where
dermal bones support the epidermis, they
are called osteoderms. (6.13)
Birds
-The integument of birds reflects some
reptilian ancestry and some new
developments of the class. Scales are
present on the legs and feet of most birds,
and the bill is covered in a tough skin that is
highly keratinized. The remaining skin is
relatively thin. The dermis of bird skin,
especially near the feather follicles, is richly
supplied with blood vessels, sensory nerve
endings, and smooth muscles. During the
breeding season, the dermis in the breast of
some birds becomes increasingly
vascularized, forming a brood patch in
which warm blood can come into close
association with incubated eggs. (6.14) -
derived originally from scales, so that scales
and feathers are homologous
- function in flight (flight feathers) as well as
temperature regulation (contour feathers)
- basic structure of feather calamus, rachis
and vane, which are derived from a feather
follicle. The vane is composed of barbs that
help to hold the shape of the feather and can
be put back into place during preening.
Birds are not always completely covered in
feathers - instead, feathers usually grow
along tracts called pterylae, and bare spots
are called aptera Some feathers are
modified to perform different functions. -
down feathers are softer feathers because
they lack all the barbs of flight feathers -
bristles and filoplumes are specially
modified feathers that are used in catching
prey (e.g., bristles around the bill of
swallows and flycatchers) and display
(filoplumes of grouse) Evolution of
Feathers When we think of feathers, we
think of their roles in flight, but they likely
had other functions when they first arose.
One view is that feathers, or their scaly
predecessors, played a role in surface
insulation. Surface insulation, of course,
holds heat in or shields the body from
taking up excess heat. Either may have been
the initial advantage of feathers. Surface
insulation would have interfered with the
absorption of environmental heat, a
disadvantage if the ancestors of birds were
ectothermic. However, many species of
ectothermic lizards have enlarged surface
scales.
Mammals
-As in other vertebrates, the two main layers
of the mammalian skin are epidermis and
dermis, which join and interface through
the basement membrane. Beneath lies the
hypodermis, or superficial fascia, composed
of connective tissue and fat. The epidermis
may be locally specialized as hair, nails, or
glands. Epithelial cells of the epidermis are
keratinocytes and belong to the keratinizing
system that forms the dead, superficial
cornified layer of the skin. The surface
keratinized cells are continually exfoliated
and replaced by cells arising primarily from
the deepest layer of the epidermis, the
stratum basale. Cells within the basale
divide mitotically, producing some that
remain to maintain the population of stem
cells and others that are pushed outward.
The mammalian dermis is double layered.
The outer papillary layer pushes
fingerlike projections, termed dermal
papillae, into the overlying epidermis. The
deeper reticular layer includes irregularly
arranged fibrous connective tissue and
anchors the dermis to the underlying fascia.
Blood vessels, nerves, and smooth muscle
occupy the dermis but do not reach the
epidermis. The mammalian dermis
produces dermal bones, but these
contribute to the skull and pectoral girdle
and only rarely form dermal scales in the
skin. Hair Hairs are slender, keratinous
filaments. The base of a hair is the root. Its
remaining length constitutes the non-living
shaft. The outer surface of the shaft often
forms a scaly cuticle. Beneath this is the
hair cortex, and at its core is the hair
medulla. Evolution of Hair Fossil skin
impressions from the Middle Jurassic give
evidence for the presence then of hair, and
presumably of mammals that owned it. But
the initial adaptive value of hair remains
speculative. One view holds that hair arose
initially as surface insulation, retaining body
heat in primitive mammalian endotherms.
The presence of nasal turbinates in
synapsids, earlier from the Late Permian,
suggests an early endothermy and, hence, a
role for insulating hair. Glands Principally
there are three main types of integumentary
glands in mammals: sebaceous, eccrine, and
apocrine. Scent, sweat, and mammary
glands are derived from them. Sebaceous
glands produce an oily secretion, sebum,
which is released into hair follicles in order
to condition and help waterproof fur.
Sebaceous glands are absent from the palms
of hands and soles of feet, but they are
present, without associated hair, at the
angle of the mouth, on the penis, near the
vagina, and next to the mammary nipples.
Eccrine glands produce thin, watery
fluids, are not associated with hair follicles,
begin to function before puberty, and are
innervated mainly by cholinergic nerves.
The apocrine glands produce a viscous,
lipid-containing fluid, are associated with
hair follicles, begin to function at puberty,
and are innervated mainly by adrenergic
nerves. Their secretions primarily function
in chemical signaling. The scent glands
are derived from apocrine glands and
produce secretions that play a part in social
communication. These glands may be
located almost anywhere on the body, as on
the chin (some deer, rabbits), face (deer,
antelope, bats), temporal region
(elephants), chest and arms (many
carnivores), anal region (rodents, dogs, cats,
mustelids), belly (musk deer), back
(kangaroo rats, peccaries, camels, ground
squirrels), or legs and feet (many
ungulates). Secretions of these glands are
used to mark territory, identify the
individual, and communicate during
courtship. The mammary glands produce
milk, a watery mixture of fats,
carbohydrates, and proteins that nourishes
the young. Ectodermal mammary ridges,
within which mammary glands form, are
located along the ventrolateral side of the
embryo. The number of mammary glands
varies among species. Release of milk to a
suckling is lactation.
SPECIALIZATIONS OF THE
INTEGUMENT
Nails, Claws, Hooves
-Nails are plates of tightly compacted,
cornified epithelial cells on the surface of
fingers and toes; thus, they are products of
the keratinizing system of the skin. The
nail matrix forms a new nail at the nail
base by pushing the existing nail forward to
replace that which is worn or broken at the
free edge. Nails protect the tips of digits
from inadvertent mechanical injury. They
also help stabilize the skin at the tips of the
fingers and toes, so that on the opposite side
the skin can establish a secure friction grip
on objects grasped. Claws, or talons, are
curved, laterally compressed keratinized
projections from the tips of digits. They are
seen in some amphibians and in most birds,
reptiles, and mammals. Hooves are
enlarged keratinized plates on the tips of the
ungulate digits. The horse hoof consists of
the hoof wall, sole, and the frog (figure
6.25). The hoof wall is U-shaped and opens
at the heel, a derivative of the integument. It
consists of a keratinized stratum externa
( tectorium), a thin, shiny surface layer; the
stratum medium, thicker and also
keratinized and permeated with coiled,
tubular channels; and an inner stratum
internum (lamellatum), a highly and
regularly laminated, infolded layer that
interdigitates with the dermis ( corium)
beneath. The hoof wall grows out from its
base, the germinal region (matrix cells), not
from the underlying dermis, at about 6 mm
per month, taking 9 to 12 months overall for
the toe to renew.
Horns and Antlers
-“Horned” lizards have processes extending
from behind the head that look like horns
but are specialized, pointed epidermal
scales. Mammals, dinosaurs, and extinct
turtles are the only vertebrates with true
horns or antlers. In horns, the associated
integument produces a tough, cornified
sheath that fits over the bony core that is
never branched. In antlers, the overlying
living skin (called “velvet”) apparently
shapes and provides vascular supply to the
growing bone. Eventually the velvet falls
away to unsheath the bare bone, the actual
material of the finished antlers that is
branched. True antlers occur only in
members of the Cervidae (e.g., deer, elk, and
moose). Typically, only males have antlers,
which are branched and shed annually.
There are notable exceptions. Among
caribou, both sexes have seasonal antlers. In
deer, the antler usually consists of a main
beam, from which branch shorter tines, or
points. In yearling bucks, antlers are
usually no more than prongs or spikes that
may be forked. The number of tines tends to
increase with age, although not exactly. In
old age, antlers may even be deformed. In
caribou and especially in moose, the main
antler beam is compressed and palmate, or
shovel-like, with a number of points
projecting from the rim. The annual cycle
of antler growth and loss in the white-
tail deer, for example, is under hormonal
control. In the spring, increasing length of
daylight stimulates the pituitary gland at the
base of the brain to release hormones that
stimulate antlers to sprout from sites on the
skull bones. By late spring, the growing
antlers are covered by velvet. By fall,
hormones produced by the testes inhibit the
pituitary, and the velvet dries. By thrashing
and rubbing, the deer wipes the velvet off to
expose the fully formed, but now dead, bone
of the antlers. Males use their antlers during
clashes with other males to maintain access
to reproductively receptive females.
Following this brief mating season, further
hormonal changes lead to a weakening of
the antler at its base where it attaches to the
living bone of the skull. The antlers break
off, and for a short time during winter, deer
are without antlers.
Among mammals, true horns are found
among members of the family Bovidae (e.g.,
cattle, antelope, sheep, goats, bison, and
wildebeests). Commonly, horns occur in
both males and females, are retained year-
round, and continue to grow throughout the
life of the individual. The horn is
unbranched and formed of a bony core and
a keratinized sheath. Those of the males are
designed to withstand the forces
encountered during head-butting combat.
In large species, females usually have horns
as well, although they are not as large and
curved as in males. In small species, females
are often hornless.
Baleen
-The integument within the mouths of
mysticete whales forms plates of baleen that
act as strainers to extract krill from water
gulped in the distended mouth. Although it
is sometimes referred to as “whalebone,” the
baleen contains no bone. It is a series of
keratinized plates that arise from the
integument. During its formation, groups of
dermal papillae extend and lengthen
outward, carrying the overlying epidermis.
The epidermis forms a cornified layer over
the surface of these projecting papillae.
Collectively, these papillae and their
covering of epidermis constitute the plates
of frilled baleen
Scales
-Scales have many functions. Both
epidermal and dermal scales are hard, so
when they receive mechanical insult and
surface abrasion, they prevent damage to
soft tissues beneath. The density of scales
also makes them a barrier against invasion
of foreign pathogens, and they retard water
loss from the body. In sharks and other
fishes, scales dampen the boundary layer
turbulence to increase swimming efficiency.
Some reptiles regulate the amount of
surface heat they absorb by turning their
bodies toward or away from the sun. This
determines whether the sun rays are
deflected off the full face of the scale or
shine under the lifted posterior edge of the
scale to reach the thin epidermis beneath.
Epidermal scales are the major component
of the skin of reptiles. They are also present
in birds along their legs, and in some
mammals, such as the beaver, they cover the
tail.
Dermal Armor
-Dermal bone forms the armor of
ostracoderm and placoderm fishes. Being a
product of the dermis, dermal bone finds its
way into alliances with a great variety of
structures. Dermal bone supports the scales
of bony fishes but tends to be lost in
tetrapods. It is absent in the skin of birds
and most mammals. Exceptions have been
noted earlier, namely, in the fossil mammal
Glyptodon and in the skin of the living
armadillo. However, selected dermal bones
take up residence in the fish skull and
pectoral girdle and have persisted into
modern groups of vertebrates. Most dermal
bones of the skull and shoulder girdle all
began phylogenetically in the skin and later
sank inward to become parts of the
skeleton. This sharing of available parts
between systems reveals again the
remodeling character of evolution.

Mucus
-Mucus produced by the skin serves several
functions. In aquatic vertebrates, it inhibits
entrance of pathogens and may even have
some slight antibacterial action. In
terrestrial amphibians, mucus keeps the
integument moist, allowing it to function in
gas exchange. Although cutaneous
respiration is prominent in amphibians, it
occurs in other vertebrates as well. For
example, many turtles rely on cutaneous gas
exchange as they hibernate submerged in
ice-covered ponds during the winter. Their
shells are too thick, of course, to allow
significant gas exchange, but exposed areas
of skin around the cloaca offer a suitable
opportunity. Sea snakes may depend on
cutaneous respiration for up to 30% of their
oxygen uptake. Similarly, fishes such as the
plaice, European eel, and mudskipper may
depend on some cutaneous gas exchange to
meet their metabolic requirements. Mucus
is also involved in aquatic locomotion. As a
surface coat, it smoothes the irregularities
and rough surface features on the epidermis
to reduce the friction met by a vertebrate
swimming through relatively viscous water.
Color
-Skin color results from complex
interactions among physical, chemical, and
structural properties of the integument.
Changes in blood supply can redden the
skin, as in blushing. The differential
scattering of light, referred to as Tyndall
scattering, is the basis for much color in
nature. This is the phenomenon that makes
the clear-day sky appear blue. In birds, air-
filled cavities within feather barbs take
advantage of this scattering phenomenon to
produce the blue feathers of kingfishers,
blue jays, bluebirds, and indigo buntings.
Many black, brown, red, orange, and yellow
colors result from pigments that produce
color by selective light reflection.
Interference phenomena are responsible for
iridescent colors. As light is reflected from
materials with different refractive indices,
interference between different wavelengths
of light produces iridescent colors. In many
birds, iridescent colors result from
interference of light reflected off the tiny
barbs and barbules of the feathers. - Many
of the pigments producing colors by this
variety of physical phenomena are
synthesized by and held in specialized
chromatophores. . Most chromatophores
arise from embryonic neural crests and can
take up residence almost anywhere within
the body. It is not uncommon to find them
associated with the walls of the digestive
tract, within the mesenteries, or around the
reproductive organs. Their function at these
remote sites is not resolved, but they are
thought to protect deep cell layers from
penetrating solar radiation. On the basis of
form, composition, and function, four
groups of chromatophores are currently
recognized. The most well-known of these is
the melanophore that contains the
pigment melanin. Melanosomes house
these melanin granules that intercept
sunlight striking the surface of an animal to
prevent penetration of harmful radiation.
They, of course, also add color to the
integument that may camouflage an animal,
making it less detectable, or brighten a part
that contributes to a behavioral display.
There are two types of melanophores. The
dermal melanophore is a broad, flat cell
that changes color rapidly and is found only
in ectotherms. The epidermal
melanophore is a thin, elongated cell
prominent in endotherms but present in all
vertebrates. By contributing melanosomes,
it adds color to keratinocytes, hair, and
feathers. The iridophore, which contains
light-reflecting, crystalline guanine
platelets, is a second type of chromatophore.
It is found in ectothermic vertebrates and in
the iris of the eye of some birds. And the
erythrophore, so called because of its red
pigments. In addition, a few
chromatophores contain several of these
pigments but are not classified. Sunlight can
influence physiological changes in
chromatophore activity. Increased exposure
stimulates increased production of pigment
granules, resulting in darker skin over a
period of days. In some vertebrates, the
response is more immediate. Some fishes
and lizards can change their colors almost
instantly. The true chameleon, for example,
can change colors to match its environment,
at least if the background is light brown to
dark green. Some fishes, such as the
flounder, can change not only their color but
also their color pattern to resemble the
background. This physiological adjustment
of color to background is mediated by the
endocrine system and involves
redistribution of pigment granules within
the chromatophores. It was once thought
that chromatophores themselves changed
shape, sending out cytoplasmic pseudopods.
Now it appears that color changes are not
based on changes in cell shape.