Exoskeleton

Last updated
The discarded exoskeleton (exuviae) of dragonfly nymph Dragonfly-nymph-exoskeleton.jpg
The discarded exoskeleton (exuviae) of dragonfly nymph
Exoskeleton of cicada attached to a Tridax procumbens Exoskeleton fly 1.jpg
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, have both an endoskeleton and an exoskeleton.

Contents

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]

Exoskeletons contain chitin; the addition of calcium carbonate makes them harder and stronger.[ citation needed ] 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. [3] 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. [4] 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

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. [5] 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[ ambiguous ] 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. [5] 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. [6]

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. BoredEncrustedShell.JPG
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 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. [7] Alternatively, exceptional preservation may result in chitin being mineralized, as in the Burgess Shale, [8] 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. [7] 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. [9]

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 [7] while others, such as Cloudina , had a calcified exoskeleton. [10] Some Cloudina shells even show evidence of predation, in the form of borings. [10]

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. 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. [11] 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. [4]

Some Precambrian (Ediacaran) organisms produced tough but non-mineralized outer shells, [7] while others, such as Cloudina, had a calcified exoskeleton, [10] 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. [12] 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. [11]

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. [4] 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. [13] A recently discovered [14] 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]

See also

Related Research Articles

Skeleton body part that forms the supporting structure of an organism

The skeleton is the body part that forms the supporting structure of an organism. It can also be seen as the bony frame work of the body which provides support, shape and protection to the soft tissues and delicate organs in animals. There are several different skeletal types: the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside the body, the hydroskeleton, a flexible skeleton supported by fluid pressure, and the cytoskeleton present in the cytoplasm of all cells, including bacteria, and archaea. The term comes from Greek σκελετός (skeletós), meaning 'dried up'.

The cloudinids, an early metazoan family containing the genera Acuticocloudina, Cloudina and Conotubus, lived in the late Ediacaran period about 550 million years ago. 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 remains unknown. The name Cloudina honors the 20th-century geologist and paleontologist Preston Cloud.

Sclerite hardened body part

A sclerite is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly to the hardened parts of arthropod exoskeletons and the internal spicules of invertebrates such as certain sponges and soft corals. In paleontology, a scleritome is the complete set of sclerites of an organism, often all that is known from fossil invertebrates.

Aragonite carbonate mineral

Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, CaCO3 (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including precipitation from marine and freshwater environments.

<i>Kimberella</i> Genus of molluscs

Kimberella is an extinct genus of bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity with this group is contentious.

Biomineralization process by which living organisms produce minerals

Biomineralization, or 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; all six taxonomic kingdoms contain members that are able to form minerals, and over 60 different minerals have been identified in organisms. Examples include silicates in algae and diatoms, carbonates in invertebrates, and calcium phosphates and carbonates in vertebrates. These minerals often form structural features such as sea shells and the bone in mammals and birds. Organisms have been producing mineralised skeletons for the past 550 million years. Ca carbonates and Ca phosphates are usually crystalline, but silica organisms (sponges, diatoms...) are always non crystalline minerals. Other examples include copper, iron and gold deposits involving bacteria. Biologically-formed minerals often have special uses such as magnetic sensors in magnetotactic bacteria (Fe3O4), gravity sensing devices (CaCO3, CaSO4, BaSO4) and iron storage and mobilization (Fe2O3•H2O in the protein ferritin).

Halkieriid Family of molluscs

The halkieriids are a group of fossil organisms from the Lower to Middle Cambrian. Their eponymous genus is Halkieria, which has been found on almost every continent in Lower to Mid Cambrian deposits, forming a large component of the small shelly fossil assemblages. The best known species is Halkieria evangelista, from the North Greenland Sirius Passet Lagerstätte, in which complete specimens were collected on an expedition in 1989. The fossils were described by Simon Conway Morris and John Peel in a short paper in 1990 in the journal Nature. Later a more thorough description was undertaken in 1995 in the journal Philosophical Transactions of the Royal Society of London and wider evolutionary implications were posed.

Namacalathus is a problematic metazoan fossil occurring in the latest Ediacaran. The first, and only described species, N. hermanastes, was first described in 2000 from the Nama Group of central and southern Namibia.

Namapoikia rietoogensis is among the earliest known animals to produce a calcareous skeleton. Known from the Ediacaran period, before the Cambrian explosion of calcifying animals, the long-lived organism grew up to a metre in diameter and resembles a colonial sponge. It was an encruster, filling vertical fissures in the reefs in which it originally grew.

The Chancelloriids are an extinct family of animal common in sediments from the Early Cambrian to the early Late Cambrian. Many of these fossils consists only of spines and other fragments, and it is not certain that they belong to the same type of organism. Other specimens appear to be more complete and to represent sessile, bag-like organisms with a soft skin armored with star-shaped calcareous sclerites from which radiate sharp spines.

Sinotubulites is a genus of small, tube-shaped shelly fossils from the Ediacaran period. It is often found in association with Cloudina.

Marine invertebrates

Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.

Cambrian substrate revolution

The "Cambrian substrate revolution" or "Agronomic revolution", evidenced in trace fossils, is the diversification of animal burrowing during the early Cambrian period.

Evidence suggesting that a mass extinction occurred at the end of the Ediacaran period, 542 million years ago, includes:

Mollusc shell Exoskeleton of an animal in the phylum Mollusca

The molluscshell 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. Not all shelled molluscs live in the sea; many live on the land and in freshwater.

The small shelly fauna, small shelly fossils (SSF), or early skeletal fossils (ESF) 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 "small shelly fauna" or "small shelly fossils". Almost all are from earlier rocks than more familiar fossils such as trilobites. Since most SSFs were preserved by being covered quickly with phosphate and this method of preservation is mainly limited to the Late Ediacaran and Early Cambrian periods, the animals that made them may actually have arisen earlier and persisted after this time span.

The Cambrian explosion or Cambrian radiation was an event approximately 541 million years ago in the Cambrian period when most major animal phyla appeared in the fossil record. It lasted for about 13 – 25 million years and resulted in the divergence of most modern metazoan phyla. The event was accompanied by major diversification of other organisms.

Nama Group lithostratigraphic unit

The Nama Group is a 125,000 square kilometres (48,000 sq mi) megaregional Vendian to Cambrian group of stratigraphic sequences deposited in the Nama foreland basin in central and southern Namibia. The Nama Basin is a peripheral foreland basin, and the Nama Group was deposited in two early basins, the Zaris and Witputs, to the north, while the South African Vanrhynsdorp Group was deposited in the southern third. The Nama Group is made of fluvial and shallow-water marine sediments, both siliciclastic and carbonate. La Tinta Group in Argentina is considered equivalent to Nama Group.

Shell growth in estuaries

Shell growth in estuaries is an aspect of marine biology that has attracted a number of scientific research studies. Many groups of marine organisms produce calcified exoskeletons, commonly known as shells, hard calcium carbonate structures which the organisms rely on for various specialized structural and defensive purposes. The rate at which these shells form is greatly influenced by physical and chemical characteristics of the water in which these organisms live. Estuaries are dynamic habitats which expose their inhabitants to a wide array of rapidly changing physical conditions, exaggerating the differences in physical and chemical properties of the water.

Marine biogenic calcification is the process by which marine organisms such as oysters and clams form calcium carbonate. Seawater is full of dissolved compounds, ions and nutrients that organisms can utilize for energy and, in the case of calcification, to build shells and outer structures. Calcifying organisms in the ocean include molluscs, foraminifera, coccolithophores, crustaceans, echinoderms such as sea urchins, and corals. The shells and skeletons produced from calcification have important functions for the physiology and ecology of the organisms that create them.

References

  1. "exoskeleton". Online Etymology Dictionary . Archived from the original on 2013-04-20.
  2. 1 2 3 S. Bengtson (2004). "Early skeletal fossils" (PDF)|chapter-format= requires |chapter-url= (help). In J. H. Lipps; B. M. Waggoner (eds.). Neoproterozoic–Cambrian Biological Revolutions. Paleontological Society Papers. 10. pp. 67–78. Archived from the original|archive-url= requires |url= (help) on 2008-10-03.
  3. 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.
  4. 1 2 3 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.
  5. 1 2 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.
  6. 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.
  7. 1 2 3 4 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. 286. London: Geological Society. pp. 157–179. Bibcode:2007GSLSP.286..157F. doi:10.1144/SP286.12. ISBN   978-1-86239-233-5. OCLC   191881597.
  8. 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.
  9. Richard Cowen (2004). History of Life (4th ed.). Wiley-Blackwell. ISBN   978-1-4051-1756-2.
  10. 1 2 3 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. doi:10.1669/0883-1351(2003)018<0454:BICSCP>2.0.CO;2.
  11. 1 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.
  12. 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.
  13. 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.
  14. 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.