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The cuticle forms the major part of the
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
of the
Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, o ...
a. It includes most of the material of the exoskeleton of the insects,
Crustacea Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
,
Arachnida Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaro ...
, and
Myriapoda Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, a ...
.


Morphology

In arthropods, the
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
, the external "skin", or "shell", is the product of a single layer of ectodermal epithelium. That layer is attached to the external or
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
surface of the deepest layer, the non-cellular internal membrane of the integument. That non-cellular membrane is called the basement membrane. The layer of epithelium on the basement membrane produces the
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
, which begins as a tough, flexible layer of chitin. Such thin, flexible chitin is the major structural part of the integument where flexibility is necessary, such as in bodily parts that must stretch to contain accumulated liquids, or that form joints between rigid parts of the exoskeleton. In other parts of the cuticle the function of the integument demands more rigid materials, such as armoured regions or the biting parts of the jaws, or where the exoskeleton forms the tubular limbs of most Arthropoda. To achieve such rigidity the outer chitin layer of the cuticle is impregnated, thickened, and reinforced with harder, more brittle materials such as
sclerotin Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which qui ...
ised proteins or
calcite Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratc ...
. This main chitinous layer of the cuticle is called the ''procuticle''. The procuticle in most land Arthropoda, particularly insects, is covered externally with a thin, waxy, water-resistant outer layer containing no chitin. That outer layer is the ''epicuticle'', and it is much thinner than the procuticle. The chitinous procuticle is formed of an outer ''exocuticle'' and the inner ''endocuticle'', and between the ''exocuticle'' and ''endocuticle'' there may be another layer called ''mesocuticle'' which has distinctive staining properties. The tough and flexible endocuticle is a laminated structure of layers of interwoven fibrous chitin and protein molecules, while the exocuticle is the layer in which any major thickening, armouring and
biomineralization Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often to harden or stiffen existing tissues. Such tissues are called mineralized tissues. It is an extremely widespread phenomenon; ...
occurs. Biomineralization with calcite is particularly common in
Crustacea Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
, whereas
sclerotization Sclerotization is a biochemical process that produces the rigid shell of sclerotin that comprises an insect's chitinous exoskeleton. It is prominent in the thicker, armored parts of insects and arachnids, especially in the biting mouthparts and scl ...
particularly occurs in insects. The exocuticle is greatly reduced in many soft-bodied insects, especially in the larval stages such as
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Symph ...
s and the larvae of parasitoidal Hymenoptera. Hardened plates in the exoskeleton are called sclerites. Sclerites may be simple protective armour, but also may form mechanical components of the exoskeleton, such as in the legs, joints, fins or wings. In the typical body segment of an insect or many other Arthropoda, there are four principal regions. The dorsal region is the
tergum A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'ma ...
; if the tergum bears any sclerites, those are called '' tergites''. The ventral region is called the sternum, which commonly bears
sternites The sternum (pl. "sterna") is the ventral portion of a segment of an arthropod thorax or abdomen. In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the ...
. The two lateral regions are called the pleura (singular pleurum) and any sclerites they bear are called pleurites. Within
entomology Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arac ...
, the term ''
glabrous Glabrousness (from the Latin ''glaber'' meaning "bald", "hairless", "shaved", "smooth") is the technical term for a lack of hair, down, setae, trichomes or other such covering. A glabrous surface may be a natural characteristic of all or part of ...
'' is used to refer to those parts of an insect's body lacking in setae (bristles) or
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
.


Chemical composition

Chemically, chitin is a long-chain polymer of a
N-acetylglucosamine ''N''-Acetylglucosamine (GlcNAc) is an amide derivative of the monosaccharide glucose. It is a secondary amide between glucosamine and acetic acid. It is significant in several biological systems. It is part of a biopolymer in the bacteria ...
, which is a derivative of glucose. The polymer bonds between the glucose units are β(1→4) links, the same as in
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
. In its unmodified form, chitin is translucent, pliable, resilient and tough. In
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, o ...
s and other organisms however, it generally is a component of a complex matrix of materials. It practically always is associated with protein molecules that often are in a more or less
sclerotised Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which qu ...
state, stiffened or hardened by cross-linking and by linkage to other molecules in the matrix. In some groups of animals, most conspicuously the
Crustacea Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
, the matrix is greatly enriched with, or even dominated by, hard minerals, usually
calcite Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratc ...
or similar
carbonate A carbonate is a salt of carbonic acid (H2CO3), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word ''carbonate'' may also refer to a carbonate ester, an organic compound containing the carbonate g ...
s that form much of the exoskeleton. In some organisms the mineral content may exceed 95%. The role of the chitin and proteins in such structures is more than just holding the crystals together; the crystal structure itself is so affected as to prevent the propagation of cracks under stress, leading to remarkable strength. The process of formation of such mineral-rich matrices is called
biomineralization Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often to harden or stiffen existing tissues. Such tissues are called mineralized tissues. It is an extremely widespread phenomenon; ...
.Campbell, N. A. (1996) ''Biology'' (4th edition) Benjamin Cummings, New Work. p.69 The difference between the unmodified and modified forms of chitinous arthropodan exoskeletons can be seen by comparing the body wall of say a bee larva, in which modification is minimal, to any armoured species of
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
, or the fangs of a spider. In both those examples there is heavy modification by sclerotisation. Again, contrasting strongly with both unmodified organic material such as largely pure chitin, and with sclerotised chitin and proteins, consider the integument of a heavily armoured crab, in which there is a very high degree of modification by biomineralization.


Moulting

As mentioned, the integument, and in particular the cuticle, is the product of the single layer of columnar or cuboidal epithelial cells attached to the basement membrane. The cuticle provides muscular support and acts as a protective shield as the insect develops, but it is not in itself cellular, so once established it cannot grow and offers little scope for maintenance, renewal or increase in size as the animal grows. Therefore, the animal periodically sheds the external part of the cuticle in a process called moulting or ecdysis. As the time for moulting approaches, as much as possible of the exocuticle material is internally digested by enzymatic action and reabsorbed through the epithelial layer. As its base gets dissolved away, the old cuticle begins to separate from the epidermis in a process called
apolysis Apolysis ( grc, ἀπόλυσις "discharge, lit. absolution") is the separation of the cuticle from the epidermis in arthropods and related groups (Ecdysozoa). Since the cuticle of these animals is also the skeletal support of the body and is in ...
. Early in the process of apolysis the epithelial cells release enzymatic moulting fluid between the old cuticle and the epidermis. The enzymes partly digest the endocuticle and the epidermis absorbs the digested material for the animal to assimilate. Much of that digested material is re-used to build the new cuticle. Once the new cuticle has formed sufficiently, the animal splits the remaining parts of the old integument along built-in lines of weakness and sheds them in the visible process of ecdysis, generally shedding and discarding the epicuticle and the reduced exocuticle, though some species carry them along for camouflage or protection. The shed portions are called the
exuvia In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have moulted. The exuviae of an animal can be important to biologists as they can often be ...
e. The animal then expands its body by swallowing liquid or gas and in the process it stretches the new integument to its proper size and shape. The new integument still is soft and usually is pale, and it is said to be teneral or callow. It then undergoes a hardening and pigmentation process that might take anything from several minutes to several days, depending on the nature of the animal and the circumstances.Gene Kritsky. (2002). ''A Survey of Entomology''. iUniverse. . File:Bienenwabe mit Eiern und Brut 5 larva.png, Honeybee larvae have flexible but delicate unsclerotised cuticles. File:Birgus latro 2.jpg, This fully-grown robber crab has tough fabric forming its joints, delicate biomineralized cuticle over its sensory antennae, optic-quality over its eyes, and strong, calcite-reinforced chitin armouring its body and legs; its pincers can break into coconuts File:Kaldari Phidippus johnsoni male defense cropped.jpg, The fangs in spiders' chelicerae are so sclerotised as to be greatly hardened and darkened


References

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Cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...