Exoskeleton

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exoskeleton.

The discarded exoskeleton (exuviae) of dragonfly nymph

Exoskeleton of cicada attached to a Tridax procumbens

An exoskeleton (from Greek έξω, éxō "outer" and σκελετός, skeletós "skeleton"[1])
is the external skeleton that supports and protects an animal's body, in contrast
to the internal skeleton (endoskeleton) of, for example, a human. In usage, some of
the larger kinds of exoskeletons are known as "shells". Examples of animals with
exoskeletons include insects such as grasshoppers and cockroaches, and crustaceans
such as crabs and lobsters, as well as the shells of certain sponges and the
various groups of shelled molluscs, including those of snails, clams, tusk shells,
chitons and nautilus. Some animals, such as the tortoise and turtle (shell page),
have both an endoskeleton and an exoskeleton.

Contents
1 Role
2 Growth
3 Paleontological significance
4 Evolution
5 See also
6 References
7 External links
Role
Exoskeletons contain rigid and resistant components that fulfill a set of
functional roles in many animals 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.[2]

Arthropod exoskeletons contain chitin; the addition of calcium carbonate makes them
harder and stronger, at the price of increased weight.[3] Ingrowths of the
arthropod exoskeleton known as apodemes serve as attachment sites for muscles.
These structures are composed of chitin and are approximately six times stronger
and twice the stiffness of vertebrate tendons. Similar to tendons, apodemes can
stretch to store elastic energy for jumping, notably in locusts.[4] Calcium
carbonates constitute the shells of molluscs, brachiopods, and some tube-building
polychaete worms. Silica forms the exoskeleton in the microscopic diatoms and
radiolaria. One species of mollusc, the scaly-foot gastropod, even makes use of the
iron sulfides greigite and pyrite.

Some organisms, such as some foraminifera, agglutinate exoskeletons by sticking

grains of sand and shell to their exterior. Contrary to a common misconception,
echinoderms do not possess an exoskeleton, as their test is always contained within
a layer of living tissue.

Exoskeletons have evolved independently many times; 18 lineages evolved calcified

exoskeletons alone.[5] Further, other lineages have produced tough outer coatings
analogous to an exoskeleton, such as some mammals. This coating is constructed from
bone in the armadillo, and hair in the pangolin. The armor of reptiles like turtles
and dinosaurs like Ankylosaurs is constructed of bone; crocodiles have bony scutes
and horny scales.
Growth
Main article: Ecdysis
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 snails, bivalves and other molluscans. A true exoskeleton, like that
found in arthropods, must be shed (moulted) when it is outgrown.[6] 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 typically stay in a den or burrow for
this time,[citation needed] as it is quite vulnerable during this period. Once at
least partially set, the organism will plump itself up to try to expand the
exoskeleton.[ambiguous] The new exoskeleton is still capable of growing to some
degree, however.[citation needed] Animals of the order arthropoda, like lizards,
amphibians, and many other animals that shed their skin, are indeterminate growers.
[1] Animals that are indeterminate growers grow in size continually throughout
their life because, in this case, their exoskeleton is always being replaced.
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, such as Azadirachtin.[7]

Paleontological significance

Borings in exoskeletons can provide evidence of animal behavior. In this case,

boring sponges attacked this hard clam shell after the death of the clam, producing
the trace fossil Entobia.
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 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.[8]
Alternatively, exceptional preservation may result in chitin being mineralized, as
in the Burgess Shale,[9] or transformed to the resistant polymer keratin, 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 are usually preserved, such as the shells of molluscs. 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.[8] The most significant limitation is
that, although there are 30-plus phyla 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.[10]

Mineralized skeletons first appear in the fossil record shortly before the base of
the Cambrian period, 550 million years ago. 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) organisms produced tough outer shells[8]
while others, such as Cloudina, had a calcified exoskeleton.[11] Some Cloudina
shells even show evidence of predation, in the form of borings.[11]

Evolution
Part of a series related to
Biomineralization

General
Exoskeletons (shells)
Arthropod exoskeletoncuticleBrachiopod shellCephalopod shell cirrate
shellcuttlebonegladiusLorica Choanoflagellate loricaMollusc shell nacrechiton
shellgastropod shellProtist shell diatom frustuleforaminifera testtestate
amoebaeSeashell echinoderm stereomsmall shelly faunascaly-foot snail shellestuary
shellsSponge spiculeTest
Endoskeletons (bones)
Teeth, scales, tusks etc
Geologic forms
Other
vte
Further information: Small shelly fauna
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. It is known, however, that in a very short course of time, just before
the Cambrian period, exoskeletons made of various materials – silica, calcium
phosphate, calcite, aragonite, and even glued-together mineral flakes – sprang up
in a range of different environments.[12] 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.[5]

Some Precambrian (Ediacaran) organisms produced tough but non-mineralized outer

shells,[8] while others, such as Cloudina, had a calcified exoskeleton,[11] but
mineralized skeletons did not become common until the beginning of the Cambrian
period, with the rise of the "small shelly fauna". 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.[13] Most other shell-forming organisms appear during the
Cambrian period, with the Bryozoans being the only calcifying phylum to appear
later, in the Ordovician. 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 proteins and polysaccharides required for the shell's composite structure, not
in the precipitation of the mineral components.[2] Skeletonization 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 skeletonization was likewise a response to increased pressure from
predators.[12]

Ocean chemistry may also control which mineral shells are constructed of. Calcium
carbonate has two forms, the stable calcite, and the metastable aragonite, 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 molluscs, 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.[5] 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.
[14] A recently discovered[15] modern gastropod Chrysomallon squamiferum that lives
near deep-sea hydrothermal vents 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 and greigite,
which had never previously been found in any metazoan but whose ingredients are
emitted in large quantities by the vents.[2]

File:Cicada exoskeleton - tokyo area - aug 15 2021.webm

Exoskeleton of a cicada
See also
Spiracle – small openings in the exoskeleton that allow insects to breathe
Hydrostatic skeleton
Endoskeleton
Powered exoskeleton
Osteoderm
References
"exoskeleton". Online Etymology Dictionary. Archived from the original on 2013-04-
20.
S. Bengtson (2004). "Early skeletal fossils" (PDF). In J. H. Lipps; B. M. Waggoner
(eds.). Neoproterozoic–Cambrian Biological Revolutions. Paleontological Society
Papers. 10. pp. 67–78. Archived from the original (PDF) on 2008-10-03.
Nedin, C. (1999). "Anomalocaris predation on nonmineralized and mineralized
trilobites". Geology. 27 (11): 987–990. Bibcode:1999Geo....27..987N.
doi:10.1130/0091-7613(1999)027<0987:APONAM>2.3.CO;2.
H. C. Bennet-Clark (1975). "The energetics of the jump of the locust, Schistocerca
gregaria" (PDF). Journal of Experimental Biology. 63 (1): 53–83. PMID 1159370.
Susannah M. Porter (2007). "Seawater chemistry and early carbonate
biomineralization". Science. 316 (5829): 1302. Bibcode:2007Sci...316.1302P.
doi:10.1126/science.1137284. PMID 17540895.
John Ewer (2005-10-11). "How the Ecdysozoan Changed Its Coat". PLOS Biology. 3
(10): e349. doi:10.1371/journal.pbio.0030349. PMC 1250302. PMID 16207077.
Gemma E. Veitch; Edith Beckmann; Brenda J. Burke; Alistair Boyer; Sarah L. Maslen;
Steven V. Ley (2007). "Synthesis of Azadirachtin: A Long but Successful Journey".
Angewandte Chemie International Edition. 46 (40): 7629–32.
doi:10.1002/anie.200703027. PMID 17665403.
M. A. Fedonkin; A. Simonetta; A. Y. Ivantsov (2007). "New data on Kimberella, the
Vendian mollusk-like organism (White sea region, Russia): palaeoecological and
evolutionary implications". In Patricia Vickers-Rich & Patricia (ed.). The Rise and
Fall of the Ediacaran Biota. Geological Society of London Special Publications.
286. London: Geological Society. pp. 157–179. Bibcode:2007GSLSP.286..157F.
doi:10.1144/SP286.12. ISBN 978-1-86239-233-5. OCLC 191881597.
Nicholas J. Butterfield (2003). "Exceptional fossil preservation and the Cambrian
Explosion". Integrative and Comparative Biology. 43 (1): 166–177.
doi:10.1093/icb/43.1.166. PMID 21680421.
Richard Cowen (2004). History of Life (4th ed.). Wiley-Blackwell. ISBN 978-1-4051-
1756-2.
Hong Hua; Brian R. Pratt; Lu-yi Zhang (2003). "Borings in Cloudina shells: complex
predator-prey dynamics in the terminal Neoproterozoic". PALAIOS. 18 (4–5): 454–459.
Bibcode:2003Palai..18..454H. doi:10.1669/0883-1351(2003)018<0454:BICSCP>2.0.CO;2.
J. Dzik (2007). "The Verdun Syndrome: simultaneous origin of protective armor and
infaunal shelters at the Precambrian–Cambrian transition" (PDF). In Patricia
Vickers-Rich & Patricia (ed.). The Rise and Fall of the Ediacaran Biota. Geological
Society, London, Special Publications. 286. London: Geological Society. pp. 405–
414. Bibcode:2007GSLSP.286..405D. CiteSeerX 10.1.1.693.9187. doi:10.1144/SP286.30.
ISBN 978-1-86239-233-5. OCLC 191881597. Archived (PDF) from the original on 2008-
10-03.
J. Dzik (1994). "Evolution of 'small shelly fossils' assemblages of the early
Paleozoic". Acta Palaeontologica Polonica. 39 (3): 27–313. Archived from the
original on 2008-12-05.
Wolfgang Kiessling; Martin Aberhan; Loïc Villier (2008). "Phanerozoic trends in
skeletal mineralogy driven by mass extinctions". Nature Geoscience. 1 (8): 527–530.
Bibcode:2008NatGe...1..527K. doi:10.1038/ngeo251.
Anders Warén; Stefan Bengtson; Shana K. Goffredi; Cindy L. Van Dover (2003). "A
hot-vent gastropod with iron sulfide dermal sclerites". Science. 302 (5647): 1007.
doi:10.1126/science.1087696. PMID 14605361.
External links
Look up exoskeleton in Wiktionary, the free dictionary.
Authority control Edit this at Wikidata
Categories: Animal anatomyBiomechanicsSkeletons
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