1398:. 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
1491:
1463:
1451:
1139:. Below a certain degree of deformation changes of shape or dimension of the cuticle are elastic and the original shape returns after the stress is removed. Beyond that level of deformation, non-reversible, plastic deformation occurs until finally the cuticle cracks or splits. Generally, the less sclerotised the cuticle, the greater the deformation required to damage the cuticle irreversibly. On the other hand, the more heavily the cuticle is armoured, the greater the stress required to deform it harmfully.
65:
40:
959:
1322:
185:
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767:
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1364:
1305:, 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.
1293:. 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
1333:
812:
1462:
1314:
28:
1407:
After the old cuticle is shed, the arthropod typically pumps up its body (for example, by air or water intake) to allow the new cuticle to expand to a larger size: the process of hardening by dehydration of the cuticle then takes place. The new integument still is soft and usually is pale, and it is
1160:
has rigid sclerites on each body segment. Supple chitin holds the sclerites together and connects the segments flexibly. Similar chitin connects the joints in the legs. Sclerotised tubular leg segments house the leg muscles, their nerves and attachments, leaving room for the passage of blood to and
1381:
The chemical and physical nature of the arthropod exoskeleton limits its ability to stretch or change shape as the animal grows. In some special cases, such as the abdomens of termite queens and honeypot ants means that continuous growth of arthropods is not possible. Therefore, growth is periodic
1215:
of an adult fly is covered with light sclerites connected by joints of membranous cuticle. In some beetles most of the joints are so tightly connected, that the body is practically in an armoured, rigid box. However, in most
Arthropoda the bodily tagmata are so connected and jointed with flexible
1439:
and life cycle roles totally different from those of the mature animals. Secondly, often a major injury in one phase, such as the loss of a leg from an insect nymph, or a claw from a young crab, can be repaired after one or two stages of ecdysis. Similarly, delicate parts that need periodic
1220:
or the larvae of mosquitoes are very mobile indeed. In addition, the limbs of arthropods are jointed, so characteristically that the very name "Arthropoda" literally means "jointed legs" in reflection of the fact. The internal surface of the exoskeleton is often infolded, forming a set of
1203:, and the tagmata are adapted to different functions in a given arthropod body. For example, tagmata of insects include the head, which is a fused capsule, the thorax as nearly a fixed capsule, and the abdomen usually divided into a series of articulating segments. Each segment has
966:
108:. Generally the exoskeleton will have thickened areas in which the chitin is reinforced or stiffened by materials such as minerals or hardened proteins. This happens in parts of the body where there is a need for rigidity or elasticity. Typically the mineral crystals, mainly
159:
of a tortoise or the cranium of a vertebrate, the exoskeleton has little ability to grow or change its form once it has matured. Except in special cases, whenever the animal needs to grow, it moults, shedding the old skin after growing a new skin from beneath.
1065:
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
118:. The crystals and fibres interpenetrate and reinforce each other, the minerals supplying the hardness and resistance to compression, while the chitin supplies the tensile strength. Biomineralization occurs mainly in crustaceans. In
1695:
1490:
1042:. By varying the types of interaction between the proteins and chitins, the insect metabolism produces regions of exoskeleton that differ in their wet and dry behaviour, their colour and their mechanical properties.
1300:
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
1120:
The two layers of the cuticle have different properties. The outer layer is where most of the thickening, biomineralization and sclerotisation takes place, and its material tends to be strong under compressive
1394:. Moulting is a complex process that is invariably dangerous for the arthropod involved. Before the old exoskeleton is shed, the cuticle separates from the epidermis through a process called
1359:, the abdomens of the workers that hold the sugar solution grow vastly, but only the unsclerotised cuticle can stretch, leaving the unstretched sclerites as dark islands on the clear abdomen
1129:, it does so by cracking. The inner layer is not as highly sclerotised, and is correspondingly softer but tougher; it resists tensile stresses but is liable to failure under compression.
1023:. The relative abundance of these two main components varies from approximately 50/50 to 80/20 chitin protein, with softer parts of the exoskeleton having a higher proportion of chitin.
1702:
1273:
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
1450:
1112:
further impregnate the cuticle with mineral salts, above all calcium carbonate, which can make up to 40% of the cuticle. The armoured product commonly has great mechanical strength.
1416:. 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.
1317:
Mature queen of a termite colony, showing how the unsclerotised cuticle stretches between the dark sclerites that failed to stretch as the abdomen grew to accommodate her ovaries
798:
1172:, 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
529:
1419:
Although the process of ecdysis is metabolically risky and expensive, it does have some advantages. For one thing it permits a complex development cycle of
1207:
according to its requirements for external rigidity; for example, in the larva of some flies, there are none at all and the exoskeleton is effectively all
1225:
that serve for the attachment of muscles, and functionally amounting to endoskeletal components. They are highly complex in some groups, particularly in
1132:
This combination is especially effective in resisting predation, as predators tend to exert compression on the outer layer, and tension on the inner.
1751:
Li, Ling; Ortiz, Christine (2014). "Pervasive nanoscale deformation twinning as a catalyst for efficient energy dissipation in a bioceramic armour".
1168:
Hardened plates in the exoskeleton are called sclerites. Sclerites may be simple protective armour, but also may form mechanical components of the
791:
1277:
state, stiffened or hardened by cross-linking and by linkage to other molecules in the matrix. In some groups of animals, most conspicuously the
1038:
chemicals crosslink protein molecules or anchor them to surrounding molecules such as chitins. Part of the effect is to make the tanned material
1814:
1565:
1199:
The arthropod exoskeleton is divided into different functional units, each comprising a series of grouped segments; such a group is called a
1527:
1371:
larva is hardly recognisable as a crab, but each time it sheds its cuticle it remodels itself, eventually taking on its final crab form
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718:
1262:, which is a derivative of glucose. The polymer bonds between the glucose units are β(1→4) links, the same as in
1589:
1238:
1007:, which is in turn secreted by the epithelial cells in the epidermis, which begins as a tough, flexible layer of
712:
371:
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1843:
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within a matrix of silk-like and globular proteins, of which the best-known is the rubbery protein called
999:
is a multi-layered external barrier that, especially in terrestrial arthropods, acts as a barrier against
683:
508:
1216:
cuticle and muscles that they have at least some freedom of movement, and many such animals, such as the
126:, the main reinforcing materials are various proteins hardened by linking the fibres in processes called
1838:
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64:
1480:, optic-quality over its eyes, and strong, calcite-reinforced chitin armouring its body and legs; its
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barely recognisable as a crab, radically changes its form when it undergoes ecdysis as it matures
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A typical arthropod exoskeleton is a multi-layered structure with four functional regions:
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Lepidoptera, Moths and
Butterflies: Morphology, Physiology, and Development : Teilband
965:
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The fangs in spiders' chelicerae are so sclerotised as to be greatly hardened and darkened
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1200:
1026:
The cuticle is soft when first secreted, but it soon hardens as required, in a process of
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Ghost crab, showing a variety of integument types in its exoskeleton, with transparent
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Its degree of sclerotisation or mineralisation determines how the cuticle responds to
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1281:, the matrix is greatly enriched with, or even dominated by, hard minerals, usually
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1321:
1242:
is used to refer to those parts of an insect's body lacking in setae (bristles) or
1082:. The exocuticle is greatly reduced in many soft-bodied insects, especially in the
743:
420:
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215:
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In its unmodified form, chitin is translucent, pliable, resilient and tough. In
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in which young animals may be totally different from older phases, such as the
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replacement, such as the outer surfaces of the eye lenses of spiders, or the
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and concentrated into a period of time when the exoskeleton is shed, called
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112:, are deposited among the chitin and protein molecules in a process called
1100:
In addition to the chitinous-proteinaceous composite of the cuticle, many
17:
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1344:, and tough chitin fabric in the joints and the bristles on the legs
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27:
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Honeybee larvae have flexible but delicate unsclerotised cuticles.
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occurs. Biomineralization with calcite is particularly common in
1003:. The strength of the exoskeleton is provided by the underlying
462:
401:
1125:, though weaker under tension. When a rigid region fails under
839:
32:
1444:
of caterpillars, can be shed, making way for new structures.
1435:, such as maggots of flies. Such larval stages commonly have
1030:. The process is poorly understood, but it involves forms of
57:, but has flexible lateral areas to allow for expansion when
1683:
10.1130/0091-7613(1999)027<0987:APONAM>2.3.CO;2
1015:, consisting of two main portions: fibrous chains of alpha-
1552:
Kristensen, Niels P.; Georges, Chauvin (1 December 2003).
1661:
predation on nonmineralized and mineralized trilobites",
1178:; if the tergum bears any sclerites, those are called
1623:(3 ed.). Oxford: Blackwell Publishing. pp.
1616:
1196:and any sclerites they bear are called pleurites.
1390:, which is under the control of a hormone called
1340:over the eyes, strong biomineralization over the
1789:(4th edition) Benjamin Cummings, New Work. p.69
1045:The chitinous procuticle is formed of an outer
1472:has tough fabric forming its joints, delicate
792:
8:
1652:
1650:
1648:
1646:
1644:
799:
785:
167:
1594:. Springer Science & Business Media.
1192:. The two lateral regions are called the
1519:
1446:
1427:of crustaceans, the nymphs of say, the
175:
35:: Chitin reinforced with sclerotisation
1615:Gullan, P. J.; P. S. Cranston (2005).
1619:The Insects: An Outline of Entomology
1011:. Arthropod cuticle is a biological
132:and the hardened proteins are called
7:
975:Dog Day Cicada moulting in Ohio USA.
86:are covered with a tough, resilient
1254:Chemically, chitin is a long-chain
1184:. The ventral region is called the
155:In either case, in contrast to the
1560:. Walter de Gruyter. p. 484.
1061:there may be another layer called
25:
49:scorpion Has heavily sclerotised
1696:"external morphology of Insects"
1588:Capinera, John L. (2008-08-11).
1489:
1461:
1449:
766:
765:
183:
530:microbial calcite precipitation
69:The cuticles of some armoured
1:
1508:Glossary of arthropod cuticle
490:marine biogenic calcification
148:form the hardened plates or
719:Biomineralising polychaetes
485:amorphous calcium carbonate
171:Part of a series related to
152:of a typical body segment.
1860:
1591:Encyclopedia of Entomology
1374:
751:Burgess Shale preservation
1476:cuticle over its sensory
1194:pleura (singular pleurum)
818:: Cuticle and epidermis;
713:Cupriavidus metallidurans
969:Time series photos of a
434:Teeth, scales, tusks etc
1785:Campbell, N. A. (1996)
1484:can break into coconuts
1188:, which commonly bears
1078:particularly occurs in
495:calcareous nannofossils
291:Choanoflagellate lorica
1807:A Survey of Entomology
1805:Gene Kritsky. (2002).
1372:
1360:
1345:
1329:
1318:
1289:that form much of the
1165:
976:
962:
955:
822:: Epicuticle detail .
684:Magnetotactic bacteria
509:oolitic aragonite sand
367:scaly-foot snail shell
78:
61:
36:
1528:"NC State University"
1366:
1351:
1335:
1324:
1316:
1157:Scutigera coleoptrata
1150:
1116:Mechanical properties
968:
961:
814:
164:Microscopic structure
67:
42:
30:
1250:Chemical composition
850:: Inner epicuticle.
846:: Outer epicuticle;
1675:1999Geo....27..987N
1657:Nedin, C. (1999), "
1431:, or the larvae of
1260:N-acetylglucosamine
411:Vertebrate skeleton
201:Mineralized tissues
1373:
1361:
1346:
1330:
1319:
1221:structures called
1166:
1090:and the larvae of
1053:, and between the
1013:composite material
977:
963:
956:
575:diatomaceous earth
541:Great Calcite Belt
458:Scale microfossils
451:otolithic membrane
362:small shelly fauna
335:echinoderm stereom
211:Biocrystallization
144:, and the lateral
79:
62:
37:
1834:Arthropod anatomy
1815:978-0-595-22143-1
1727:"Insect Glossary"
1567:978-3-11-016210-3
1468:This fully-grown
1338:biomineralization
1295:biomineralization
1068:biomineralization
995:. Of these, the
993:basement membrane
888:Basement membrane
809:
808:
739:permineralization
724:Mineral nutrients
649:Mineral evolution
318:foraminifera test
177:Biomineralization
115:biomineralization
110:calcium carbonate
16:(Redirected from
1851:
1818:
1803:
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1765:10.1038/nmat3920
1753:Nature Materials
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1701:. Archived from
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1605:
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1539:
1530:. Archived from
1524:
1493:
1465:
1453:
1442:urticating hairs
1108:and the extinct
954:: Gland opening.
834:: Cement layer;
801:
794:
787:
774:
769:
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689:Magnetoreception
669:Ballast minerals
264:Cephalopod shell
259:Brachiopod shell
206:Remineralisation
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1669:(11): 987–990,
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1425:nauplius larvae
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1153:house centipede
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1086:stages such as
805:
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313:diatom frustule
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1759:(5): 501–507.
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1375:Main article:
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1076:sclerotization
1049:and the inner
1028:sclerotization
896:epidermal cell
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570:siliceous ooze
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557:Silicification
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379:Sponge spicule
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1809:. iUniverse.
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1795:0-8053-1957-3
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1708:on 2011-07-19
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1634:1-4051-1113-5
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1601:9781402062421
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1534:on 2008-09-06
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1421:metamorphosis
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734:Fossilization
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729:Microbial mat
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674:Magnetofossil
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514:aragonite sea
512:
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477:Calcification
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398:Endoskeletons
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136:. The dorsal
135:
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19:
1839:Biomechanics
1806:
1801:
1786:
1781:
1756:
1752:
1746:
1734:. Retrieved
1730:
1721:
1710:. Retrieved
1703:the original
1690:
1666:
1662:
1659:Anomalocaris
1658:
1618:
1610:
1590:
1583:
1571:. Retrieved
1557:
1554:"Integument"
1547:
1536:. Retrieved
1532:the original
1522:
1418:
1413:
1409:
1406:
1399:
1380:
1354:honeypot ant
1299:
1268:
1253:
1237:
1231:
1198:
1179:
1173:
1167:
1155:
1143:Segmentation
1134:
1131:
1119:
1099:
1092:parasitoidal
1088:caterpillars
1062:
1058:
1054:
1050:
1046:
1044:
1025:
978:
970:
951:
947:
939:
931:
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899:
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851:
847:
843:
835:
831:
823:
819:
815:
744:petrifaction
711:
699:
694:Microfossils
441:Limpet teeth
421:Ossification
416:Bone mineral
350:chiton shell
246:
235:Exoskeletons
216:Biointerface
154:
133:
127:
113:
99:
93:
87:
81:
80:
73:are rigidly
44:
43:This female
1736:21 February
1470:robber crab
1408:said to be
1291:exoskeleton
1285:or similar
1275:sclerotised
1236:, the term
1170:exoskeleton
1137:deformation
1102:crustaceans
1095:Hymenoptera
1063:mesocuticle
1059:endocuticle
1051:endocuticle
1040:hydrophobic
1001:desiccation
864:Endocuticle
679:Magnetosome
622:phosphorite
588:Other forms
536:calcite sea
303:coccosphere
247:exoskeleton
101:exoskeleton
75:sclerotised
53:, tail and
1828:Categories
1731:E-Fauna BC
1712:2011-03-20
1573:10 January
1538:2008-07-16
1514:References
1437:ecological
1369:Zoea-stage
1326:Crab larva
1287:carbonates
1271:arthropods
1234:entomology
1209:membranous
1110:trilobites
1074:, whereas
1055:exocuticle
1047:exocuticle
1034:in which
1005:procuticle
997:epicuticle
985:procuticle
981:epicuticle
904:Pore canal
872:Procuticle
856:Exocuticle
828:Epicuticle
274:cuttlebone
243:Arthropod
140:, ventral
89:integument
83:Arthropods
18:Epicuticle
1279:Crustacea
1264:cellulose
1227:Crustacea
1218:Chilopoda
1205:sclerites
1190:sternites
1161:from the
1106:myriapods
1072:Crustacea
989:epidermis
920:Trichogen
912:Glandular
880:Epidermis
700:engrailed
617:Phosphate
610:oil shale
504:Aragonite
308:coccolith
150:sclerites
134:sclerotin
124:arachnids
1773:24681646
1502:See also
1478:antennae
1396:apolysis
1392:ecdysone
1384:moulting
1357:repletes
1309:Moulting
1239:glabrous
1223:apodemes
1181:tergites
1163:hemocoel
1123:stresses
1036:phenolic
950:: Hair;
944:Sensilia
928:Tormogen
772:Category
653:In soil
605:alginite
595:Bone bed
330:Seashell
237:(shells)
157:carapace
46:Pandinus
1787:Biology
1671:Bibcode
1663:Geology
1482:pincers
1429:Odonata
1410:teneral
1401:exuviae
1388:ecdysis
1377:Ecdysis
1342:pincers
1283:calcite
1256:polymer
1232:Within
1213:abdomen
1186:sternum
1104:, some
1080:insects
1032:tanning
1021:resilin
972:Tibicen
842:layer;
642:Related
600:Kerogen
525:Calcite
446:Otolith
279:gladius
252:cuticle
221:Biofilm
194:General
142:sternum
120:insects
95:cuticle
71:beetles
1813:
1793:
1771:
1631:
1598:
1564:
1414:callow
1303:beetle
1244:scales
1211:; the
1175:tergum
1127:stress
1084:larval
1017:chitin
1009:chitin
930:cell;
922:cell;
914:cell;
770:
629:Pyrena
286:Lorica
146:pleura
138:tergum
106:chitin
59:gravid
55:dorsum
51:chelae
1706:(PDF)
1699:(PDF)
1625:22–24
1367:This
1258:of a
1201:tagma
936:Nerve
707:Druse
402:bones
345:nacre
1811:ISBN
1791:ISBN
1769:PMID
1738:2017
1629:ISBN
1596:ISBN
1575:2013
1562:ISBN
1151:The
1057:and
991:and
702:gene
463:Tusk
384:Test
122:and
1761:doi
1679:doi
1412:or
1386:or
1352:In
868:2+3
840:Wax
104:of
98:or
33:ant
1830::
1767:.
1757:13
1755:.
1729:.
1677:,
1667:27
1665:,
1643:^
1627:.
1556:.
1404:.
1297:.
1266:.
1246:.
1229:.
1097:.
987:,
983:,
952:13
948:12
946:;
942::
940:11
938:;
934::
932:10
926::
918::
910::
906:;
902::
900:6a
898:;
894::
890:;
886::
882:;
878::
874:;
870::
866:;
862::
858:;
854::
848:1d
844:1c
838::
836:1b
832:1a
830:;
826::
92:,
1817:.
1775:.
1763::
1740:.
1715:.
1681::
1673::
1637:.
1604:.
1577:.
1541:.
924:9
916:8
908:7
892:6
884:5
876:4
860:3
852:2
824:1
820:B
816:A
800:e
793:t
786:v
404:)
400:(
20:)
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