Exoskeleton
Encyclopedia
An exoskeleton is the external skeleton
Skeleton
The skeleton is the body part that forms the supporting structure of an organism. There are two different skeletal types: the exoskeleton, which is the stable outer shell of an organism, and the endoskeleton, which forms the support structure inside the body.In a figurative sense, skeleton can...

 that supports and protects an animal's body, in contrast to the internal skeleton (endoskeleton
Endoskeleton
An endoskeleton is an internal support structure of an animal, composed of mineralized tissue. Endoskeleton develops within the skin or in the deeper body tissues. The vertebrate is basically an endoskeleton made up of two types of tissues . During early embryonic development the endoskeleton is...

) of, for example, a human
Human skeleton
The human skeleton consists of both fused and individual bones supported and supplemented by ligaments, tendons, muscles and cartilage. It serves as a scaffold which supports organs, anchors muscles, and protects organs such as the brain, lungs and heart....

. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s such as grasshopper
Grasshopper
The grasshopper is an insect of the suborder Caelifera in the order Orthoptera. To distinguish it from bush crickets or katydids, it is sometimes referred to as the short-horned grasshopper...

s and cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...

es, and crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...

s such as crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

s and lobster
Lobster
Clawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...

s. The shells
Mollusc shell
The mollusc shell is typically a calcareous exoskeleton which encloses, supports and protects the soft parts of an animal in the phylum Mollusca, which includes snails, clams, tusk shells, and several other classes...

 of the various groups of shelled mollusks, including those of snail
Snail
Snail is a common name applied to most of the members of the molluscan class Gastropoda that have coiled shells in the adult stage. When the word is used in its most general sense, it includes sea snails, land snails and freshwater snails. The word snail without any qualifier is however more often...

s, clam
Clam
The word "clam" can be applied to freshwater mussels, and other freshwater bivalves, as well as marine bivalves.In the United States, "clam" can be used in several different ways: one, as a general term covering all bivalve molluscs...

s, tusk shell
Tusk shell
The tusk shells or scaphopods are a class of shelled marine mollusks. The scientific name of this class is Scaphopoda, meaning "shovel-footed". Shells of species within this class range from about 0.5 to 15 cm in length...

s, chiton
Chiton
Chitons are small to large, primitive marine molluscs in the class Polyplacophora.There are 900 to 1,000 extant species of chitons in the class, which was formerly known as Amphineura....

s and nautilus
Nautilus
Nautilus is the common name of marine creatures of cephalopod family Nautilidae, the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in two genera, the type of which is the genus Nautilus...

, are also exoskeletons.

Mineralized exoskeletons first appeared in the fossil record about , and their evolution is considered by some to have played a role in the subsequent Cambrian explosion
Cambrian explosion
The Cambrian explosion or Cambrian radiation was the relatively rapid appearance, around , of most major phyla, as demonstrated in the fossil record, accompanied by major diversification of other organisms, including animals, phytoplankton, and calcimicrobes...

 of animals.

Some animals, such as the tortoise
Tortoise
Tortoises are a family of land-dwelling reptiles of the order of turtles . Like their marine cousins, the sea turtles, tortoises are shielded from predators by a shell. The top part of the shell is the carapace, the underside is the plastron, and the two are connected by the bridge. The tortoise...

, have both an endoskeleton and an exoskeleton.

Role of the exoskeleton

Exoskeletons contain rigid and resistant components that fulfil a set of functional roles including protection, excretion, sensing, support, feeding and acting as a barrier against desiccation in terrestrial organisms. Exoskeletons have a role in defense from pests and predators, support, and in providing an attachment framework for musculature.

Exoskeletons contain chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...

 and when calcium carbonate is added, the exoskeleton grows in strength and hardness.

Ingrowths of the arthropod exoskeleton known as apodemes serve as attachment sites for muscle
Muscle
Muscle is a contractile tissue of animals and is derived from the mesodermal layer of embryonic germ cells. Muscle cells contain contractile filaments that move past each other and change the size of the cell. They are classified as skeletal, cardiac, or smooth muscles. Their function is to...

s. These structures are composed of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...

, and are approximately 6 times as strong and twice as stiff as vertebrate tendon
Tendon
A tendon is a tough band of fibrous connective tissue that usually connects muscle to bone and is capable of withstanding tension. Tendons are similar to ligaments and fasciae as they are all made of collagen except that ligaments join one bone to another bone, and fasciae connect muscles to other...

s. Similar to tendons, apodemes can stretch to store elastic energy for jumping, notably in locust
Locust
Locusts are the swarming phase of short-horned grasshoppers of the family Acrididae. These are species that can breed rapidly under suitable conditions and subsequently become gregarious and migratory...

s.

Diversity

Many produce exoskeletons, which are composed of a range of materials. Bone, cartilage, or dentine is used in the Ostracoderm
Ostracoderm
Ostracoderms are any of several groups of extinct, primitive, jawless fishes that were covered in an armor of bony plates. They belong to the taxon Ostracodermi, and their fossils are found in the Ordovician and Devonian Period strata of North America and Europe...

 fish and turtle
Turtle
Turtles are reptiles of the order Testudines , characterised by a special bony or cartilaginous shell developed from their ribs that acts as a shield...

s. Chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...

 forms the exoskeleton in arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s including insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s, arachnid
Arachnid
Arachnids are a class of joint-legged invertebrate animals in the subphylum Chelicerata. All arachnids have eight legs, although in some species the front pair may convert to a sensory function. The term is derived from the Greek words , meaning "spider".Almost all extant arachnids are terrestrial...

s such as spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...

s, crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...

s such as crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

s and lobster
Lobster
Clawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...

s (see arthropod exoskeleton
Arthropod exoskeleton
Arthropods are covered with a tough or resistant exoskeleton, which may be mineralised or constructed of a tough polymer such as chitin. This external skeleton is moulted as the organism grows.- Microscopic structure :...

), and in some fungi
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...

 and bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

. Calcium carbonates constitute the shells of molluscs
Mollusca
The Mollusca , common name molluscs or mollusksSpelled mollusks in the USA, see reasons given in Rosenberg's ; for the spelling mollusc see the reasons given by , is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest...

 (see Mollusc shell
Mollusc shell
The mollusc shell is typically a calcareous exoskeleton which encloses, supports and protects the soft parts of an animal in the phylum Mollusca, which includes snails, clams, tusk shells, and several other classes...

), brachiopod
Brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...

s, and some tube-building polychaete
Polychaete
The Polychaeta or polychaetes are a class of annelid worms, generally marine. Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made of chitin. Indeed, polychaetes are sometimes referred to as bristle worms. More than 10,000...

 worms. Silica
Silicon dioxide
The chemical compound silicon dioxide, also known as silica , is an oxide of silicon with the chemical formula '. It has been known for its hardness since antiquity...

 forms the exoskeleton in the microscopic diatom
Diatom
Diatoms are a major group of algae, and are one of the most common types of phytoplankton. Most diatoms are unicellular, although they can exist as colonies in the shape of filaments or ribbons , fans , zigzags , or stellate colonies . Diatoms are producers within the food chain...

s and radiolaria
Radiolarian
Radiolarians are amoeboid protozoa that produce intricate mineral skeletons, typically with a central capsule dividing the cell into inner and outer portions, called endoplasm and ectoplasm. They are found as zooplankton throughout the ocean, and their skeletal remains cover large portions of the...

.

Some organisms, such as some foraminifera
Foraminifera
The Foraminifera , or forams for short, are a large group of amoeboid protists which are among the commonest plankton species. They have reticulating pseudopods, fine strands of cytoplasm that branch and merge to form a dynamic net...

, agglutinate exoskeletons by sticking grains of sand and shell to their exterior. Contrary to a common misconception, echinoderm
Echinoderm
Echinoderms are a phylum of marine animals. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone....

s do not possess an exoskeleton, as their test
Test (biology)
A test is a term used to refer to the shell of sea urchins, and also the shell of certain microorganisms, such as testate foraminifera and testate amoebae....

 is always contained within a layer of living tissue.

Exoskeletons have evolved independently many times; 18 lineages evolved calcified
Calcification
Calcification is the process in which calcium salts build up in soft tissue, causing it to harden. Calcifications may be classified on whether there is mineral balance or not, and the location of the calcification.-Causes:...

 exoskeletons alone. Further, other lineages have produced tough outer coatings analogous to an exoskeleton, such as some mammals – (constructed from bone in the armadillo
Armadillo
Armadillos are New World placental mammals, known for having a leathery armor shell. Dasypodidae is the only surviving family in the order Cingulata, part of the superorder Xenarthra along with the anteaters and sloths. The word armadillo is Spanish for "little armored one"...

, and hair in the pangolin
Pangolin
A pangolin , also scaly anteater or Trenggiling, is a mammal of the order Pholidota. There is only one extant family and one genus of pangolins, comprising eight species. There are also a number of extinct taxa. Pangolins have large keratin scales covering their skin and are the only mammals with...

) – and reptiles (turtle and Ankylosaur
Ankylosauria
Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. They are first known to have appeared in the early Jurassic Period of...

 armor are constructed of bone; crocodiles have bony scute
Scute
A scute or scutum is a bony external plate or scale, as on the shell of a turtle, the skin of crocodilians, the feet of some birds or the anterior portion of the mesonotum in insects.-Properties:...

s and horn
Horn (anatomy)
A horn is a pointed projection of the skin on the head of various animals, consisting of a covering of horn surrounding a core of living bone. True horns are found mainly among the ruminant artiodactyls, in the families Antilocapridae and Bovidae...

y scales).

Growth in an exoskeleton

Since exoskeletons are rigid, they present some limits to growth. Organisms with open shells can grow by adding new material to the aperture of their shell, as is the case in snail
Snail
Snail is a common name applied to most of the members of the molluscan class Gastropoda that have coiled shells in the adult stage. When the word is used in its most general sense, it includes sea snails, land snails and freshwater snails. The word snail without any qualifier is however more often...

s, bivalves and other mollusca
Mollusca
The Mollusca , common name molluscs or mollusksSpelled mollusks in the USA, see reasons given in Rosenberg's ; for the spelling mollusc see the reasons given by , is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest...

ns. A true exoskeleton, like that found in in arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s must be shed (moulted
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...

) when they are outgrown. A new exoskeleton is produced beneath the old one. As the old one is shed, the new skeleton is soft and pliable. The animal will pump itself up to expand the new shell to maximal size, then let it harden. When the shell has set, the empty space inside the new skeleton can be filled up as the animal eats. Failure to shed the exoskeleton once outgrown can result in the animal being suffocated within its own shell, and will stop subadults from reaching maturity, thus preventing them from reproducing. This is the mechanism behind some insect pesticides, like Azadirachtin
Azadirachtin
Azadirachtin, a chemical compound belonging to the limonoid group, is a secondary metabolite present in neem seeds. It is a highly oxidized tetranortriterpenoid which boasts a plethora of oxygen functionality, comprising an enol ether, acetal, hemiacetal, and tetra-substituted oxirane as well as a...

.

Paleontological significance

Exoskeletons, as hard parts of organisms, are greatly useful in assisting preservation of organisms, whose soft parts usually rot before they can be fossilized. Mineralized exoskeletons can be preserved "as is", as shell fragments, for example. The possession of an exoskeleton also permits a couple of other routes to fossilization. For instance, the tough layer can resist compaction, allowing a mold of the organism to be formed underneath the skeleton, which may later decay. Alternatively, exceptional preservation
Lagerstätte
A Lagerstätte is a sedimentary deposit that exhibits extraordinary fossil richness or completeness.Palaeontologists distinguish two kinds....

 may result in chitin being mineralized, as in the Burgess Shale
Burgess Shale
The Burgess Shale Formation, located in the Canadian Rockies of British Columbia, is one of the world's most celebrated fossil fields, and the best of its kind. It is famous for the exceptional preservation of the soft parts of its fossils...

, or transformed to the resistant polymer keratin
Keratin
Keratin refers to a family of fibrous structural proteins. Keratin is the key of structural material making up the outer layer of human skin. It is also the key structural component of hair and nails...

, which can resist decay and be recovered.

However our dependence on fossilized skeletons also significantly limits our understanding of evolution. Only the parts of organisms that were already mineralized
Mineralization (biology)
In biology, mineralization refers to the process where an organic substance is converted to an inorganic substance.This may also be a normal biological process which takes place during the life of an organism such as the formation of bone tissue or egg shells, largely with calcium.This term may...

 are usually preserved, such as the shells of mollusks. It helps that exoskeletons often contain "muscle scars", marks where muscles have been attached to the exoskeleton, which may allow the reconstruction of much of an organism's internal parts from its exoskeleton alone. The most significant limitation is that, although there are 30-plus phyla
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....

 of living animals, two-thirds of these phyla have never been found as fossils, because most animal species are soft-bodied and decay before they can become fossilized.

Mineralized skeletons first appear in the fossil record shortly before the base of the Cambrian period, . The evolution of a mineralized exoskeleton is seen by some as a possible driving force of the Cambrian explosion of animal life, resulting in a diversification of predatory and defensive tactics. However, some Precambrian (Ediacaran
Ediacaran
The Ediacaran Period , named after the Ediacara Hills of South Australia, is the last geological period of the Neoproterozoic Era and of the Proterozoic Eon, immediately preceding the Cambrian Period, the first period of the Paleozoic Era and of the Phanerozoic Eon...

) organisms
Ediacara biota
The Ediacara biota consisted of enigmatic tubular and frond-shaped, mostly sessile organisms which lived during the Ediacaran Period . Trace fossils of these organisms have been found worldwide, and represent the earliest known complex multicellular organisms.Simple multicellular organisms such as...

 produced tough outer shells, while others, such as Cloudina
Cloudinid
The Cloudinids, an early metazoan family containing the genus Cloudina, lived in the late Ediacaran period and became extinct at the base of the Cambrian. They formed millimetre-scale conical fossils consisting of calcareous cones nested within one another; the appearance of the organism itself...

, had a calcified exoskeleton.
Some Cloudina shells even show evidence of predation, in the form of borings.

Evolution

On the whole, the fossil record only contains mineralised exoskeletons, since these are by far the most durable. Since most lineages with exoskeletons are thought to have started out with a non-mineralised exoskeleton which they later mineralised, this makes it difficult to comment on the very early evolution of each lineage's exoskeleton. We do know that in a very short course of time just before the Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...

 period exoskeletons made of various materials – silica, calcium phosphate
Calcium phosphate
Calcium phosphate is the name given to a family of minerals containing calcium ions together with orthophosphates , metaphosphates or pyrophosphates and occasionally hydrogen or hydroxide ions ....

, calcite
Calcite
Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate . The other polymorphs are the minerals aragonite and vaterite. Aragonite will change to calcite at 380-470°C, and vaterite is even less stable.-Properties:...

, aragonite
Aragonite
Aragonite is a carbonate mineral, one of the two common, naturally occurring, crystal forms of calcium carbonate, CaCO3...

, and even glued-together mineral flakes – sprang up in a range of different environments. Most lineages adopted the form of calcium carbonate which was stable in the ocean at the time they first mineralised, and did not change from this mineral morph - even when it became the less favorable.

Some Precambrian (Ediacaran) organisms produced tough but non-mineralized outer shells, while others, such as Cloudina, had a calcified exoskeleton, but mineralized skeletons did not become common until the beginning of the Cambrian period, with the rise of the "small shelly fauna
Small shelly fauna
The small shelly fauna or small shelly fossils, abbreviated to SSF, are mineralized fossils, many only a few millimetres long, with a nearly continuous record from the latest stages of the Ediacaran to the end of the Early Cambrian period. They are very diverse, and there is no formal definition of...

". Just after the base of the Cambrian, these miniature fossils become diverse and abundant – this abruptness may be an illusion, since the chemical conditions which preserved the small shellies appeared at the same time. Most other shell forming organisms appear during the Cambrian period, with the Bryozoa
Bryozoa
The Bryozoa, also known as Ectoprocta or commonly as moss animals, are a phylum of aquatic invertebrate animals. Typically about long, they are filter feeders that sieve food particles out of the water using a retractable lophophore, a "crown" of tentacles lined with cilia...

ns being the only calcifying phylum to appear later, in the Ordovician
Ordovician
The Ordovician is a geologic period and system, the second of six of the Paleozoic Era, and covers the time between 488.3±1.7 to 443.7±1.5 million years ago . It follows the Cambrian Period and is followed by the Silurian Period...

. The sudden appearance of shells has been linked to a change in ocean chemistry which made the calcium compounds of which the shells are constructed stable enough to be precipitated into a shell. However this is unlikely to be a sufficient cause, as the main construction cost of shells is in creating the protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

s and polysaccharide
Polysaccharide
Polysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...

s required for the shell's composite structure
Composite material
Composite materials, often shortened to composites or called composition materials, are engineered or naturally occurring materials made from two or more constituent materials with significantly different physical or chemical properties which remain separate and distinct at the macroscopic or...

, not in the precipitation of the mineral components. Skeletonisation also appeared at almost exactly the same time that animals started burrowing to avoid predation, and one of the earliest exoskeletons was made of glued-together mineral flakes, suggesting that skeletonisation was likewise a response to increased pressure from predators.

Ocean chemistry may also control which mineral shells are constructed of. Calcium carbonate has two forms, the stable calcite, and the metastable
Metastability
Metastability describes the extended duration of certain equilibria acquired by complex systems when leaving their most stable state after an external action....

 aragonite
Aragonite
Aragonite is a carbonate mineral, one of the two common, naturally occurring, crystal forms of calcium carbonate, CaCO3...

, which is stable within a reasonable range of chemical environments but rapidly becomes unstable outside this range. When the oceans contain a relatively high proportion of magnesium compared to calcium, aragonite is more stable, but as the magnesium concentration drops, it becomes less stable, hence harder to incorporate into an exoskeleton, as it will tend to dissolve.

With the exception of the mollusks, whose shells often comprise both forms, most lineages use just one form of the mineral. The form used appears to reflect the seawater chemistry – thus which form was more easily precipitated – at the time that the lineage first evolved a calcified skeleton, and does not change thereafter. However, the relative abundance of calcite- and aragonite-using lineages does not reflect subsequent seawater chemistry – the magnesium/calcium ratio of the oceans appears to have a negligible impact on organisms' success, which is instead controlled mainly by how well they recover from mass extinctions. A recently-discovered modern gastropod that lives near deep-sea hydrothermal vent
Hydrothermal vent
A hydrothermal vent is a fissure in a planet's surface from which geothermally heated water issues. Hydrothermal vents are commonly found near volcanically active places, areas where tectonic plates are moving apart, ocean basins, and hotspots. Hydrothermal vents exist because the earth is both...

s illustrates the influence of both ancient and modern local chemical environments: its shell is made of aragonite, which is found in some of the earliest fossil mollusks; but it also has armor plates on the sides of its foot, and these are mineralized with the iron sulfides pyrite
Pyrite
The mineral pyrite, or iron pyrite, is an iron sulfide with the formula FeS2. This mineral's metallic luster and pale-to-normal, brass-yellow hue have earned it the nickname fool's gold because of its resemblance to gold...

 and greigite
Greigite
Greigite is an iron sulfide mineral with formula Fe3S4. It is the sulfur equivalent of the iron oxide magnetite . It was first described in 1964 for an occurrence in San Bernardino County, California, and named after the mineralogist and physical chemist Joseph W...

, which had never previously been found in any metazoan
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

 but whose ingredients are emitted in large quantities by the vents.

Artificial "exoskeletons"

Human
Human
Humans are the only living species in the Homo genus...

s have long used armor as an artificial exoskeleton for protection, especially in combat. Exoskeletal machines (also called powered exoskeleton
Powered exoskeleton
A powered exoskeleton, also known as powered armor, or exoframe, is a powered mobile machine consisting primarily of an exoskeleton-like framework worn by a person and a power supply that supplies at least part of the activation-energy for limb movement.Powered exoskeletons are designed to assist...

s) are also starting to be used for medical and industrial purposes, while powered human exoskeletons are a feature of science fiction writing, but are currently moving into prototype stage.

Orthoses
Orthotics
Orthotics is a specialty within the medical field concerned with the design, manufacture and application of orthoses. An orthosis is an orthopedic device that supports or corrects the function of a limb or the torso...

 are a limited, medical form of exoskeleton. An orthosis (plural orthoses) is a device which attaches to a limb, or the torso, to support the function or correct the shape of that limb or the spine. Orthotics
Orthotics
Orthotics is a specialty within the medical field concerned with the design, manufacture and application of orthoses. An orthosis is an orthopedic device that supports or corrects the function of a limb or the torso...

 is the field dealing with orthoses, their use, and their manufacture. An orthotist is a person who designs and fits orthoses. A prosthesis
Prosthesis
In medicine, a prosthesis, prosthetic, or prosthetic limb is an artificial device extension that replaces a missing body part. It is part of the field of biomechatronics, the science of using mechanical devices with human muscle, skeleton, and nervous systems to assist or enhance motor control...

 (plural prostheses) is a device that substitutes for a missing part of a limb. If the prosthesis is a hollow shell and self-carrying, it is exoskeletal. If internal tubes are used in the device and the cover (cosmesis
Cosmesis
Cosmesis is the preservation, restoration, or bestowing of bodily beauty. In the medical context, it usually refers to the surgical correction of a disfiguring defect, or the cosmetic improvements made by a surgeon following incisions...

) to create the outside shape is made of a soft, non-carrying material, it is endoskeletal. Prosthetics is the field that deals with prostheses, use, and their manufacture. A prosthetist
Prosthetist
A prosthetist, as defined by , is a person who measures, designs, fabricates, fits, or services a prosthesis as prescribed by a licensed physician, and who assists in the formulation of the prosthesis prescription for the replacement of external parts of the human body lost due to amputation or...

 is a person who designs and fits prostheses.

Perhaps the first animals to use a naturally-occurring "artificial exoskeleton" were the hermit crab
Hermit crab
Hermit crabs are decapod crustaceans of the superfamily Paguroidea. Most of the 1100 species possess an asymmetrical abdomen which is concealed in an empty gastropod shell that is carried around by the hermit crab.-Description:...

s, the majority of which are obliged constantly to "wear" an empty gastropod shell, in order to protect their soft abdomens.

Parenthetically, the exoskeleton has been used as an architectural model as in the St. Martin Island Light
St. Martin Island Light
St. Martin Island Light is a unique exoskeleton lighthouse that marks one of four passages between Lake Michigan and the bay of Green Bay. Constructed in 1905, this light tower is the only example in the United States of a pure exoskeletal tower on the Great Lakes. Similar designs do exist in...

. The building technique is called monocoque
Monocoque
Monocoque is a construction technique that supports structural load by using an object's external skin, as opposed to using an internal frame or truss that is then covered with a non-load-bearing skin or coachwork...

, and is used in architecture and in construction of car
Čar
Čar is a village in the municipality of Bujanovac, Serbia. According to the 2002 census, the town has a population of 296 people.-References:...

s and aeroplanes.

See also

  • Mechatronics
    Mechatronics
    Mechatronics is the combination of mechanical engineering, electronic engineering, computer engineering, software engineering, control engineering, and systems design engineering in order to design, and manufacture useful products. Mechatronics is a multidisciplinary field of engineering, that is...

  • Powered exoskeleton
    Powered exoskeleton
    A powered exoskeleton, also known as powered armor, or exoframe, is a powered mobile machine consisting primarily of an exoskeleton-like framework worn by a person and a power supply that supplies at least part of the activation-energy for limb movement.Powered exoskeletons are designed to assist...

  • Spiracle
    Spiracle
    Spiracles are openings on the surface of some animals that usually lead to respiratory systems.-Vertebrates:The spiracle is a small hole behind each eye that opens to the mouth in some fishes. In the primitive jawless fish the first gill opening immediately behind the mouth is essentially similar...

     – small openings in the exoskeleton that allow insects to breathe
  • Hydrostatic skeleton
    Hydrostatic skeleton
    A hydrostatic skeleton or hydroskeleton is a structure found in many cold-blooded organisms and soft-bodied animals consisting of a fluid-filled cavity, the coelom, surrounded by muscles. The pressure of the fluid and action of the surrounding circular and longitudinal muscles are used to change an...

  • Endoskeleton
    Endoskeleton
    An endoskeleton is an internal support structure of an animal, composed of mineralized tissue. Endoskeleton develops within the skin or in the deeper body tissues. The vertebrate is basically an endoskeleton made up of two types of tissues . During early embryonic development the endoskeleton is...


Biology


Modern technology

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