INTEGUMENT SYSTEM

SOME ROLES OF THE INTEGUMENT

● Protective Role
○ Primary role of the integument
○ Dermal armors protects the internal organs from mechanical injury
○ Glands secrete slimy or noxious substances
○ Pigments provide protective coloration
○ Serves as barrier to the rays of the sun
○ Claws, horns, spiny protuberances and needles give protection
● Exteroceptive Role
○ Protective in its most primitive state
○ Naked nerve endings are stimulated when foreign objects contact the skin
■ The only role of cutaneous receptors in agnathans
○ More complex receptors are found in other fishes and tetrapods and highly used for survival
● Respiration
○ Supplements gills and lungs of many amphibians (aquatic urodeles)
○ Plethodontid salamander rely its respiration entirely on the skin (they don’t have gills or lungs)
○ Scales and cornified epithelium are not conducive for cutaneous respiration
● Excretion
○ CO2 excretion in some aquatic amphibians
○ Sweat glands are supplementary to excrete nitrogenous wastes
○ Ammonia in fishes is easily diffused via gill epithelium and other tissues exposed to water
● Thermoregulation
○ Function of the skin of endotherms (birds and mammals)
○ Fur and feather insulate against cold
○ Sweat cools by evaporation
○ Dilation of blood vessels within the dermis increases heat loss by radiation
● Locomotion
○ Performed by adhesive pads, claws that assist in climbing, scutes that assist in slithering and feathers that provide
airfoil
○ Webbed feet for those that wade in water
○ Webbed wings in bats enable them to fly
● Maintenance of Homeostasis
○ Reservoirs of calcium and phosphate molecules in fishes
○ Cornified epithelium of tetrapods conserve water
○ Absorbs water under the influence of posterior pituitary gland in aestivating lungfishes, toads, & other craniates
● Nourishment
○ Mucus secreted in some teleosts provide nutrition to hatchlings
○ Mammary glands provide nourishment to the young
● Pheromones and Skin Coloration
○ Serves as species identification or serves as alarm
○ Elaborate colorations help during breeding season
○ Can be used as protection

GENERAL STRUCTURE AND DEVELOPMENT

Integument
● Forms the interface between the animal’s internal environment and the outside world
● Contains epithelial, connective adipose and smooth muscle tissues
● Also contains blood vessels, glands, sensory receptors, nerves and other structures
Epidermis
● Composed of a stratified epithelium which developed from embryonic ectoderm
● Rests on a basement membrane made up of delicate fibrils and overlies the dermis
● Stratum germinativum consists of 1-2 mitotically active, cuboidal cell layers located above the basement membrane
○ Cells move toward the body surface to differentiate and are eventually sloughed off

● Some epidermal cells synthesize keratin which is a water-insoluble, horny protein that may fill the cells and replace other
organelles
○ Best developed in terrestrial vertebrates in which keratin-filled cells form the stratum corneum
● Fishes and amphibians - thin layer of epidermis
● Reptiles and mammals - thick layer of epidermis
● Two types of covering:
○ Aquatic craniates - thin coat of mucus
○ Terrestrial craniates - water-impervious cornified cells
● Gland cells are epidermal in origin but invade the dermis
○ Aquatic craniates - predominantly unicellular
○ Terrestrial craniates - mostly multicellular

Dermis
● Develops from mesenchymal cells which are mostly derived from mesodermal dermatome of somites
● Basic component is collagen and elastic fibers embedded in proteoglycans and other macromolecules which holds
other component in place and provide tensile strength
● Other components: blood vessels, small nerves, and pigment cells
○ Lymphatics
○ Naked and encapsulated exteroceptors
○ Bases of multicellular glands
○ Bases of hairs or feathers and their erector muscles
● Dermis is thicker than the epidermis in mature skin and consists of two layers:
○ Stratum laxum - irregularly arranged collagen fibers
○ Stratum compactum - irregularly arranged collagen fibers

Layers of the Skin

Skin Derivatives
● Teeth, bony and horny scales, feathers and hair develop as a result of intricate interactions between the dermis and
epidermis via neural crest cell mitigation
● Glands, feathers, hair and other keratinized structures are composed of epidermal cells in which some parts invade the
underlying dermis
● Superficial bones are dermal derivatives
● Teeth and bony scales are of epidermal and dermal products
● These structures have supportive and protective functions that they are sometimes termed as integumentary skeleton
Chromatophores
● Cells that contain pigments found in all vertebrates except albinos
● Develop from neural crests and are located in the upper part of the dermis in fishes, amphibians, and reptiles
● They penetrate and are located in the epidermis of birds and mammals

Melanophores
● Star-shaped cells containing melanin
● With long, branching processes
● Melanin granules may either be blac, brown or more yellow and reddish
● Contained within cellular organelles called melanosomes
● Melanin is synthesized within melanophores of birds and mammals but most of it is transferred to feather, hair, and
other epidermal derivatives
● Melanosomes in anamniotes and reptiles may migrate into processes of the melanophores, maximizing the color or
concentrating it

Other Pigment Secreting Cells

● Iridophores or Guanophores
○ Contain crystalline chemochromes from guanine
○ Generate iridescent colors because of the diffraction of light within the stacked plates
● Leucophores
○ Utilize crystalline purines (often guanine) to reflect light
○ Produce reflective white hues
● Xanthophores
○ Contain yellow pigments composed of pteridines
● Erythrophores
○ Contain reddish pigments composed of carotenoids
● Vesicles containing pteridine and carotenoids are sometimes found in the same cell

Dermal Chromatophore Unit

Turacoverdin and Turacin

● Turacoverdin
○ Unique copper uroporphyrin pigment responsible for the bright green coloration of several birds of the
family Musophagidae, most notably the turaco
○ The only true green pigments found in birds
● Turacin
○ Naturally occurring red pigment
○ Found only in the bird family Musophagidae, the turacos

Important Roles of Skin Coloration

● Can be concealing or cryptic helping an animal hide from predators or enable a predator to stalk a prey
● Can be aposematic where it advertises the presence of dangerous, venomous or distasteful species
● For species recognition, establishment of territories, courtship and other types of communication
● Helps reptiles thermoregulate

FISH INTEGUMENT
Epidermis
● Integument of fishes is similar in its basic structure
● Epidermis is relatively thin and most cells are alive
● Surface cells are covered with microridges that increase the surface area for exchange between the animal & its
environment
● Keratin may be deposited in limited areas such as horny teeth of cyclostomes and tubercles that develop in many fishes
during mating season
Epidermal Glands
● Goblet cells - secrete only mucus
● Granular cells - secrete mucus and additional ingredients mostly of unknown functions
○ Alkaloids which are products of granular cells are not common

● Multicellular glands - not abundant

● Slimy mucus can be for protection, for nutrition, and some contain toxins

Photophores
● Light-emitting organs in deep-sea teleosts
● Upper part of the gland consists of modified mucous cells acting as magnifying lens
● Beneath the lens are light-emitting cells
● Cells below are in contact with melanophores and in blood cells which is the source of raw materials
○ Luciferase + luciferin = light
● The light is not intense and usually of many hues
● Functions for:
○ Species and sex identification
○ Lure for prey in carnivores or is a warning that it is a carnivore
○ Concealment by countershading

Epidermis of Agnathans
● Highly mitotic, multi-layered epidermis with unicellular mucous glands
● No keratinized or cornified layer
● Horny denticles on buccal funnel or teeth are the only keratinized structures

Epidermis of Cartilaginous Fishes

● Multi-layered than agnathans
● Not as much unicellular glands except in chimeras
● Localized glands:
○ Goblet cells - stingray
○ Multicellular glands - male claspers
● Photophores in the dermis are modified epidermal glands that lost connection with epidermis

Epidermis of Bony Fishes

● Top layer composed of stratified epidermal cells that do not undergo keratinization
● Lowermost layer is the basal layer
● Glands are mostly unicellular (mucous coat on skin) with few multicellular and granular glands
● Photophores for recognition or warning

Dermis
● Collagen fibers are generally more regularly arranged than of other vertebrates
● Develop in layers that spiral around the body in approximately 45o angle to the longitudinal axis with adjacent layers
perpendicular to each other
 ● Fiber arrangement strengthens the skin so that body shape is maintained during swimming
● Acts as exotendon transmitting muscular force

Bony Dermis of Fishes

● Generalized pattern:
4 = enamel or enamel-like substance
3 = dentin (denticles is 3 +4)
2 = vascular spongy bone
1 = lamellar bone

Derivatives of Primitive Dermal Bone

Dermal Scales

Dermis of Agnathans
● No dermal bone
● Thinner than epidermis
● Tough due to collagen connective tissue
● With many melanophores
● With slime glands

Dermis of Cartilaginous Fishes

● No dermal bone
● Surface denticles (placoid scales) present except in chimeras
● Thicker than the epidermis
● With melanophores (more dorsally)

Dermis of Bony Fishes

● Cosmoid and ganoid in basal actinopterygians and neopterygians (Polypterus and garpikes)
● Cycloid and ctenoid in modern fishes
AMPHIBIAN INTEGUMENT
Epidermis
● Skin of amphibian is relatively thin but epidermal cells synthesize keratin
● As it accumulates, cells die and the stratum corneum is formed which is seldom more than 1-2 layers thick to allow
cutaneous respiration
○ Desquamated periodically and is hormonally controlled
○ Not sloughed off in toads and they just continually pile up
● Highly glandular
● Glandular glands aid in survival of land-adapted anurans
● Transitional between those that spend more time and more on land
● Cornified appendages are rare

Epidermal Glands
● Mostly multicellular mucous or granular glands
● Tailed aquatic amphibians have the most number of multicellular glands
○ Secretions keep the skin moist when they are on land
● Glands on digits serves as holdfasts in trees or glands on thumb pads during breeding season serves as restraint for females

Granular Glands
● Present in toads
● Secrete irritating alkaloids which are defensive in nature or pheromones used during breeding
● Restricted to a localized area of the body

Keratin
● Aquatic urodeles have thin desiccation-impending stratum of cornified cells
● Anuran tadpoles have horny tooth like structures which is shed during metamorphosis

Dermis
● Firmly attached to underlying muscles in apodans and urodeles
● Anurans have lymph sinuses separating it from muscles
● Dermal chromatophores in other species can elicit color change
● Bony scales (osteoderms) present in head of caecilians and a few tropical toads at the back

REPTILE INTEGUMENT
Epidermis
● Represents the ultimate adaptation of craniate skin for surviving in an arid and hostile environment
● With many localized modifications: horny scales, scutes, beaks, rattles, claws, plaques, and spiny crests found on the
stratum corneum
● Granular glands are found in some region only

Granular Glands
Epidermal Scales
● Repetitious thickenings only found in amniotes
● Disposed on overlapping folds of the epidermis in squamates where thinning at the scale joints allow for movement
● Scutes are large, thin, polygonal scales

Molting

Dermis
● With dermal bone
● Turtles - carapace, plastron, and lateral bridges
○ Soft-shelled and leatherback turtles lack dermal ossification
● Crocodiles and some lizards - osteoderms in localized regions of the body (gastralia in ventrolateral abdominal wall)
● Snakes - osteoderm is absent

Dermal Ossification in Tetrapods

AVIAN INTEGUMENT
Epidermis
● Scales only present at the feet and base of the beak
● Claws are present on the toes and one or two on digits of the wings
● Glands are generally lacking except:
○ Uropygial glands at the tail base for preening
○ Oil glands at the outer ear canal of domesticated birds

Avian Oil Glands or Uropygial Gland

● Uropygial gland
○ Prominent swelling at the rump behind the pygostyle
○ Largest in aquatic birds and in domestic fowl
○ Oil which is water-repellent is used for preening
● Smaller oil glands are found in outer ear canal and sometimes in the vent area

Feathers
Feather Tracts

Development of a Feather
Mesodermal Component Epidermal Component

Dermal Papilla is formed Feather Primordium is induced

Dermal Papilla becomes vascularized Feather Follicle develops as Feather Primordium elongates

Growth Zone develops at the base of the Feather Follicle,

giving rise to the Feather Sheath

Feather Sheath splits open as the shaft elongates to full growth

Dermal Papilla dies when the feather is fully grown

and becomes pulp as the Inferior Umbilicus is formed

Dermis
● Supports feather follicles and erector, depressor, retractor and rotator muscles
● Feathers are cornified appendages first developed for insulation and is a modified scale
● No osteoderms (except in gamecocks with a spur of dermal bone at the ankle)

MAMMALIAN INTEGUMENT
Epidermis
● Stratum germinativum - basal layer
● Stratum granulosum - with keratohyalin
● Stratum corneum - highly keratinized
○ Give rise to scales (armadillos and pangolins), claws (hoofs and flat nails), and horns
○ Thickest on parts of appendages in contact with surfaces (palms, hooves, feet)
● Stratum lucidum - present in palm and sole
● Hair are cornified epidermal appendages
● Claws are present in most mammals and can be modified as nails in primates or hooves in ungulates
● Other cornified appendages include horns, pronghorn, antlers, baleen of whales and epidermal scales

Hair
 Development of a Hair Hair Life Cycle

Claws, Hoofs and Nails Bovine Horns and Pronghorns

Hair Horns Other Cornified Structures

Epidermal Glands
● Structure
○ Tubular
■ Simple
■ Branched
■ Coiled
■ Compound
○ Alveolar
■ Simple
■ Branched
■ Compound
● Mode of Elaboration
○ Merocrine
○ Apocrine
○ Holocrine
● Substance secreted
○ Mucous
○ Granular
○ Avian oil
○ Sebaceous
○ Sweat
○ Scent
○ Mammary
 Epidermal Glands Based on Structure

Epidermal Glands Based on Mode of Elaboration

Mucous Glands
● Generally absent in tetrapods except in mammals
● Lost survival value when skin became covered with cornified cells
● Synthesis of mucus will dehydrate craniates that don’t have continuous access to water

Sebaceous Glands
● Present when there are hairs and sebum is exuded into hair follicles
● Ceruminous glands secrete cerumen in outer ear
● Meibomian glands assist in lubrication of conjunctiva of the eye
● Chalazion which is an inflamed swelling on the conjunctival surface of the lid is caused by ducts of the Meibomian
glands being occluded
● Other sebaceous glands open independent of hair:
○ Lips
○ Glans penis
○ Labia minora
○ Areola

Sudoriferous or Sweat Glands

● Furry animals - feet of cat & mice, lips of rabbits & side of head of bats
● Hippopotamus - only on the ears
● Absent in pangolins, sirenians, cetaceans, and echidnas
● Ciliary glands is one example

Scent Glands
Mammary Glands
● Compound alveolar glands that develop in both sexes along the milk line
● Number and location (axillary, thoracic, abdominal, and inguinal) is dependent on the animal species
● Oxytocin is responsible for milk-let down

Dermis
● Very thick - hair follicles, erector muscle, numerous glands, connective tissue, vascular tissue, nervous tissue
● Dermal bones of armadillos
● Antlers and giraffe horns
● Superficial fascia separates the dermis from underlying muscles
● Blubber is a type of contour-shaping adipose tissue found in cetaceans to replace hair

Dermal Bones, Antlers, and Giraffe Horns