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Skin
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For skin in humans, see Human skin. For other uses, see Skin (disambiguation).
Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with
three main functions: protection, regulation, and sensation.
Fur is dense hair.[8] Primarily, fur augments the insulation the skin provides but can also serve as a
secondary sexual characteristic or as camouflage. On some animals, the skin is very hard and thick
and can be processed to create leather. Reptiles and most fish have hard protective scales on their
skin for protection, and birds have hard feathers, all made of tough beta-keratins. Amphibian skin is
not a strong barrier, especially regarding the passage of chemicals via skin, and is often subject to
osmosis and diffusive forces. For example, a frog sitting in an anesthetic solution would be sedated
quickly as the chemical diffuses through its skin. Amphibian skin plays key roles in everyday survival
and their ability to exploit a wide range of habitats and ecological conditions.[9]
On 11 January 2024, biologists reported the discovery of the oldest known skin, fossilized about 289
million years ago, and possibly the skin from an ancient reptile.[10][11]
Contents
Etymology
Structure in mammals
Epidermis
Basement membrane
Dermis
Papillary region
Reticular region
Subcutaneous tissue
Amphibians
Overview
Granular glands
Mucous glands
Development
Functions
Mechanics
Aging
See also
References
External links
Etymology
The word skin originally only referred to dressed and tanned animal hide and the usual word for human
skin was hide. Skin is a borrowing from Old Norse skinn "animal hide, fur", ultimately from the Proto-
Indo-European root *sek-, meaning "to cut" (probably a reference to the fact that in those times
animal hide was commonly cut off to be used as garment).[12]
Structure in mammals
Epidermis
Main article: Epidermis
The distribution of the blood vessels in the skin of
The epidermis is composed of the outermost the sole of the foot. (Corium – TA alternate term for
layers of the skin. It forms a protective barrier dermis – is labeled at upper right.)
over the body's surface, responsible for
keeping water in the body and preventing
pathogens from entering, and is a stratified
squamous epithelium,[13] composed of
proliferating basal and differentiated
suprabasal keratinocytes.
Stratum corneum
A diagrammatic sectional view of the skin (click on
Stratum lucidum (only in palms and soles) image to magnify). (Dermis labeled at center right.)
Stratum granulosum
Identifiers
Stratum spinosum
MeSH D012867
Stratum basale (also called the stratum
germinativum) TA98 A16.0.00.002
The epidermis contains no blood vessels, and cells in the deepest layers are nourished by diffusion
from blood capillaries extending to the upper layers of the dermis.
Basement membrane
Main article: Basement membrane
The epidermis and dermis are separated by a thin sheet of fibers called the basement membrane,
which is made through the action of both tissues. The basement membrane controls the traffic of the
cells and molecules between the dermis and epidermis but also serves, through the binding of a
variety of cytokines and growth factors, as a reservoir for their controlled release during physiological
remodeling or repair processes.[16]
Dermis
Main article: Dermis
The dermis is the layer of skin beneath the epidermis that consists of connective tissue and cushions
the body from stress and strain. The dermis provides tensile strength and elasticity to the skin through
an extracellular matrix composed of collagen fibrils, microfibrils, and elastic fibers, embedded in
hyaluronan and proteoglycans.[15] Skin proteoglycans are varied and have very specific locations.[17]
For example, hyaluronan, versican and decorin are present throughout the dermis and epidermis
extracellular matrix, whereas biglycan and perlecan are only found in the epidermis.
It harbors many mechanoreceptors (nerve endings) that provide the sense of touch and heat through
nociceptors and thermoreceptors. It also contains the hair follicles, sweat glands, sebaceous glands,
apocrine glands, lymphatic vessels and blood vessels. The blood vessels in the dermis provide
nourishment and waste removal from its own cells as well as for the epidermis.
Dermis and subcutaneous tissues are thought to contain germinative cells involved in formation of
horns, osteoderm, and other extra-skeletal apparatus in mammals.[2]
The dermis is tightly connected to the epidermis through a basement membrane and is structurally
divided into two areas: a superficial area adjacent to the epidermis, called the papillary region, and a
deep thicker area known as the reticular region.
Papillary region
The papillary region is composed of loose areolar connective tissue. This is named for its fingerlike
projections called papillae that extend toward the epidermis. The papillae provide the dermis with a
"bumpy" surface that interdigitates with the epidermis, strengthening the connection between the two
layers of skin.
Reticular region
The reticular region lies deep in the papillary region and is usually much thicker. It is composed of
dense irregular connective tissue and receives its name from the dense concentration of collagenous,
elastic, and reticular fibers that weave throughout it. These protein fibers give the dermis its properties
of strength, extensibility, and elasticity. Also located within the reticular region are the roots of the
hair, sweat glands, sebaceous glands, receptors, nails, and blood vessels.
Subcutaneous tissue
Main article: Subcutaneous tissue
The subcutaneous tissue (also hypodermis) is not part of the skin, and lies below the dermis. Its
purpose is to attach the skin to underlying bone and muscle as well as supplying it with blood vessels
and nerves. It consists of loose connective tissue and elastin. The main cell types are fibroblasts,
macrophages and adipocytes (the subcutaneous tissue contains 50% of body fat). Fat serves as
padding and insulation for the body.
Microorganisms like Staphylococcus epidermis colonize the skin surface. The density of skin flora
depends on region of the skin. The disinfected skin surface gets recolonized from bacteria residing in
the deeper areas of the hair follicle, gut and urogenital openings.
Fish
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The epidermis of fish and of most amphibians consists entirely of live cells, with only minimal
quantities of keratin in the cells of the superficial layer.[18] It is generally permeable, and in the case of
many amphibians, may actually be a major respiratory organ.[19] The dermis of bony fish typically
contains relatively little of the connective tissue found in tetrapods. [18] Instead, in most species, it is
largely replaced by solid, protective bony scales.[20] Apart from some particularly large dermal bones
that form parts of the skull, these scales are lost in tetrapods, although many reptiles do have scales
of a different kind, as do pangolins.[21] Cartilaginous fish have numerous tooth-like denticles
embedded in their skin, in place of true scales.[22]
Sweat glands and sebaceous glands are both unique to mammals, but other types of skin gland are
found in other vertebrates.[23] Fish typically have a numerous individual mucus-secreting skin cells
that aid in insulation and protection, but may also have poison glands, photophores, or cells that
produce a more watery, serous fluid. In amphibians, the mucus cells are gathered together to form
sac-like glands. Most living amphibians also possess granular glands in the skin, that secrete irritating
or toxic compounds.[24]
Although melanin is found in the skin of many species, in the reptiles, the amphibians, and fish, the
epidermis is often relatively colorless. Instead, the color of the skin is largely due to chromatophores in
the dermis, which, in addition to melanin, may contain guanine or carotenoid pigments. Many species,
such as chameleons and flounders may be able to change the color of their skin by adjusting the
relative size of their chromatophores.[24]
Amphibians
Further information: Amphibian § Skin
Overview
Amphibians possess two types of glands, mucous and granular (serous). Both of these glands are part
of the integument and thus considered cutaneous. Mucous and granular glands are both divided into
three different sections which all connect to structure the gland as a whole. The three individual parts
of the gland are the duct, the intercalary region, and lastly the alveolar gland (sac). Structurally, the
duct is derived via keratinocytes and passes through to the surface of the epidermal or outer skin
layer thus allowing external secretions of the body. The gland alveolus is a sac-shaped structure that
is found on the bottom or base region of the granular gland. The cells in this sac specialize in
secretion. Between the alveolar gland and the duct is the intercalary system which can be summed up
as a transitional region connecting the duct to the grand alveolar beneath the epidermal skin layer. In
general, granular glands are larger in size than the mucous glands, which are greater in number.[25]
G
r
a
n
u
l
a
r
g
l
a
n
d
s
Frog Gland Anatomy– A: Mucous gland (alveolus), B: Chromophore, C: Granular Gland (alveolus), D: Connective Tissue,
E: Stratum Corneum, F: Transition Zone (intercalary region), G: Epidermis (Where the duct resides), H: Dermis
Granular glands can be identified as venomous and often differ in the type of toxin as well as the
concentrations of secretions across various orders and species within the amphibians. They are
located in clusters differing in concentration depending on amphibian taxa. The toxins can be fatal to
most vertebrates or have no effect against others. These glands are alveolar meaning they structurally
have little sacs in which venom is produced and held before it is secreted upon defensive
behaviors.[25]
Structurally, the ducts of the granular gland initially maintain a cylindrical shape. When the ducts
mature and fill with fluid, the base of the ducts become swollen due to the pressure from the inside.
This causes the epidermal layer to form a pit like opening on the surface of the duct in which the inner
fluid will be secreted in an upwards fashion.[26]
The intercalary region of granular glands is more developed and mature in comparison with mucous
glands. This region resides as a ring of cells surrounding the basal portion of the duct which are
argued to have an ectodermal muscular nature due to their influence over the lumen (space inside the
tube) of the duct with dilation and constriction functions during secretions. The cells are found radially
around the duct and provide a distinct attachment site for muscle fibers around the gland's body.[26]
The gland alveolus is a sac that is divided into three specific regions/layers. The outer layer or tunica
fibrosa is composed of densely packed connective-tissue which connects with fibers from the spongy
intermediate layer where elastic fibers, as well as nerves, reside. The nerves send signals to the
muscles as well as the epithelial layers. Lastly, the epithelium or tunica propria encloses the gland.[26]
Mucous glands
Mucous glands are non-venomous and offer a different functionality for amphibians than granular.
Mucous glands cover the entire surface area of the amphibian body and specialize in keeping the body
lubricated. There are many other functions of the mucous glands such as controlling the pH,
thermoregulation, adhesive properties to the environment, anti-predator behaviors (slimy to the
grasp), chemical communication, even anti-bacterial/viral properties for protection against
pathogens.[25]
The ducts of the mucous gland appear as cylindrical vertical tubes that break through the epidermal
layer to the surface of the skin. The cells lining the inside of the ducts are oriented with their
longitudinal axis forming 90-degree angles surrounding the duct in a helical fashion.[26]
Intercalary cells react identically to those of granular glands but on a smaller scale. Among the
amphibians, there are taxa which contain a modified intercalary region (depending on the function of
the glands), yet the majority share the same structure.[26]
The alveolar or mucous glands are much more simple and only consist of an epithelium layer as well as
connective tissue which forms a cover over the gland. This gland lacks a tunica propria and appears to
have delicate and intricate fibers which pass over the gland's muscle and epithelial layers.[26]
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The epidermis of birds and reptiles is closer to that of mammals, with a layer of dead keratin-filled
cells at the surface, to help reduce water loss. A similar pattern is also seen in some of the more
terrestrial amphibians such as toads. In these animals, there is no clear differentiation of the epidermis
into distinct layers, as occurs in humans, with the change in cell type being relatively gradual. The
mammalian epidermis always possesses at least a stratum germinativum and stratum corneum, but
the other intermediate layers found in humans are not always distinguishable. Hair is a distinctive
feature of mammalian skin, while feathers are (at least among living species) similarly unique to
birds.[24]
Birds and reptiles have relatively few skin glands, although there may be a few structures for specific
purposes, such as pheromone-secreting cells in some reptiles, or the uropygial gland of most
birds.[24]
Development
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Cutaneous structures arise from the epidermis and include a variety of features such as hair, feathers,
claws and nails. During embryogenesis, the epidermis splits into two layers: the periderm (which is
lost) and the basal layer. The basal layer is a stem cell layer and through asymmetrical divisions,
becomes the source of skin cells throughout life. It is maintained as a stem cell layer through an
autocrine signal, TGF alpha, and through paracrine signaling from FGF7 (keratinocyte growth factor)
produced by the dermis below the basal cells. In mice, over-expression of these factors leads to an
overproduction of granular cells and thick skin.[27][28]
Hair and feathers are formed in a regular pattern and it is believed to be the result of a reaction-
diffusion system. This reaction-diffusion system combines an activator, Sonic hedgehog, with an
inhibitor, BMP4 or BMP2, to form clusters of cells in a regular pattern. Sonic hedgehog-expressing
epidermal cells induce the condensation of cells in the mesoderm. The clusters of mesodermal cells
signal back to the epidermis to form the appropriate structure for that position. BMP signals from the
epidermis inhibit the formation of placodes in nearby ectoderm. [citation needed]
It is believed that the mesoderm defines the pattern. The epidermis instructs the mesodermal cells to
condense and then the mesoderm instructs the epidermis of what structure to make through a series
of reciprocal inductions. Transplantation experiments involving frog and newt epidermis indicated that
the mesodermal signals are conserved between species but the epidermal response is species-
specific meaning that the mesoderm instructs the epidermis of its position and the epidermis uses this
information to make a specific structure.[29]
Functions
1. Protection: an anatomical barrier from pathogens and damage between the internal and external
environment in bodily defense. (See Skin absorption.) Langerhans cells in the skin are part of the
adaptive immune system.[3][4]
2. Sensation: contains a variety of nerve endings that jump to heat and cold, touch, pressure, vibration,
and tissue injury (see somatosensory system and haptic perception).
3. Thermoregulation: Eccrine (sweat) glands and dilated blood vessels (increased superficial perfusion)
aid heat loss, while constricted vessels greatly reduce cutaneous blood flow and conserve heat.
Erector pili muscles in mammals adjust the angle of hair shafts to change the degree of insulation
provided by hair or fur.
4. Control of evaporation: the skin provides a relatively dry and semi-impermeable barrier to reduce fluid
loss.[4]
5. Storage and synthesis: acts as a storage center for lipids and water
6. Absorption through the skin: Oxygen, nitrogen and carbon dioxide can diffuse into the epidermis in
small amounts; some animals use their skin as their sole respiration organ (in humans, the cells
comprising the outermost 0.25–0.40 mm of the skin are "almost exclusively supplied by external
oxygen", although the "contribution to total respiration is negligible")[30] Some medications are
absorbed through the skin.
7. Water resistance: The skin acts as a water resistant barrier so essential nutrients aren't washed out of
the body. The nutrients and oils that help hydrate the skin are covered by the most outer skin layer, the
epidermis. This is helped in part by the sebaceous glands that release sebum, an oily liquid. Water
itself will not cause the elimination of oils on the skin, because the oils residing in our dermis flow and
would be affected by water without the epidermis.[31]
8. Camouflage, whether the skin is naked or covered in fur, scales, or feathers, skin structures provide
protective coloration and patterns that help to conceal animals from predators or prey.[32]
Mechanics
Main article: Soft tissue
Skin is a soft tissue and exhibits key mechanical behaviors of these tissues. The most pronounced
feature is the J-curve stress strain response, in which a region of large strain and minimal stress exists
and corresponds to the microstructural straightening and reorientation of collagen fibrils.[33] In some
cases the intact skin is prestreched, like wetsuits around the diver's body, and in other cases the
intact skin is under compression. Small circular holes punched on the skin may widen or close into
ellipses, or shrink and remain circular, depending on preexisting stresses.[34]
Aging
Tissue homeostasis generally declines with age, in part because stem/progenitor cells fail to self-
renew or differentiate. Skin aging is caused in part by TGF-β by blocking the conversion of dermal
fibroblasts into fat cells which provide support. Common changes in the skin as a result of aging range
from wrinkles, discoloration, and skin laxity, but can manifest in more severe forms such as skin
malignancies.[35][36] Moreover, these factors may be worsened by sun exposure in a process known as
photoaging.[36]
See also
Moult
Skinning
References
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doi:10.1111/brv.12597 . PMID 32338826 . Ellen; Finke, Mark; Fleming, Greg (2014). "Nutrition
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External links