Difference between revisions of "Sea urchin" - New World Encyclopedia

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{{Taxobox
 
{{Taxobox
 
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| name = Sea urchin
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***Order [[Holectypoida]]
 
***Order [[Holectypoida]]
 
}}
 
}}
'''Sea urchin''' is the common name for various spiky [[echinoderm]]s within the class '''Echinoidea''', characterized by pentamerous [[radial symmetry]]; a hard calcareous shell, or test, covered with movable spines that is made of regularly arranged, tightly fused plates through which rows of slender, extensible tube feet extend; and a mouth on the lower side, facing the substratum, with a complex chewing structure composed of five jaws (Aristotle's lantern). In a more general sense of the term, sea urchin sometimes is used as the common name for all members of [[Echinoidea]]. However, more specifically, it is used to refer to the "regular echinoids" within subclass Euechinoidea that are pentaradially symmetrical and have their anus located on the aboral surface (opposite to the mouth).
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'''Sea urchin''' is the common name for various spiky [[echinoderm]]s within the class '''Echinoidea''', characterized by pentamerous [[radial symmetry]]; a hard calcareous shell, or test, made of regularly arranged, tightly fused plates through which rows of slender, extensible tube feet extend; movable spines covering the shell; and a mouth on the lower side, facing the substratum, with a complex chewing structure composed of five jaws (Aristotle's lantern).  
  
The term sea urchin, thus does not normally include the pencil urchins in subclass Perishoechinoidea, nor the "irregular echinoids" within Euechinoidea, which include the heart urchins and [[sand dollar]]s. The later tend to be characterized by a secondary [[bilateral symmetry]] and a more flattened and oval test, with very short spines, and with the anus either posterior or on the oral surface.
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In a more general sense of the term, sea urchin sometimes is used as the common name for all members of [[Echinoidea]]. However, more specifically, it is used to refer to the "regular echinoids" within subclass Euechinoidea that are pentaradially symmetrical and have their anus located on the aboral surface (opposite to the mouth). The term sea urchin, thus, does not normally include the pencil urchins in subclass Perishoechinoidea, nor the "irregular echinoids" within Euechinoidea, which include the heart urchins and [[sand dollar]]s. The later tend to be characterized by a secondary [[bilateral symmetry]] and a more flattened and oval test, with very short spines, and with the anus either posterior or on the oral surface.
  
They are found in oceans all over the world.
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Sea urchins are found in oceans all over the world. They play an important role in marine [[food chain]]s, consuming [[algae]] and various [[invertebrate]]s, and being consumed by [[crab]]s, [[sea star]]s, [[fish]]es, [[mammal]]s, and [[bird]]s. When predators such as [[sea otter]]s disappear from the ecosystem, the unchecked sea urchins can devastate their environment, making an "urchin barren."
 
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{{toc}}
Sea urchins are harvested and served as a delicacy.
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For humans, sea urchins are harvested and served as a delicacy. Their roe and gonads are frequently consumed raw, or briefly cooked, with species such as the red sea urchin ''(Strongylocentrotus francis-canus)'', green urchin ''(S. droebachiensis)'', and purple sea urchin ''(S. purpuratus)'' among those favored for their roe (Freeman 2004). They are a frequent model organism for developmental and immunological studies.
  
 
==Overview and description==
 
==Overview and description==
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[[Image:Zeeegel3.jpg|right|240px|thumb|Sea urchin test. Each white band is the location of a row of tube feet; each pair of white bands is called an ambulacrum. There are five such ambulacra; the fivefold symmetry reveals a kinship with starfish.]]
 
Sea urchins are members of the [[echinoderm|phylum Echinodermata]], which also includes [[starfish]], [[sea cucumber]]s, [[brittle star]]s, and [[crinoid]]s. Like other echinoderms they have fivefold symmetry (called [[symmetry (biology)#Pentamerism|pentamerism]]) and move by means of hundreds of tiny, transparent, adhesive "[[tube feet]]." The pentamerous symmetry is not obvious at a casual glance but is easily seen in the dried shell or test of the urchin.
 
Sea urchins are members of the [[echinoderm|phylum Echinodermata]], which also includes [[starfish]], [[sea cucumber]]s, [[brittle star]]s, and [[crinoid]]s. Like other echinoderms they have fivefold symmetry (called [[symmetry (biology)#Pentamerism|pentamerism]]) and move by means of hundreds of tiny, transparent, adhesive "[[tube feet]]." The pentamerous symmetry is not obvious at a casual glance but is easily seen in the dried shell or test of the urchin.
  
The class '''Echinoidea''' comprises sea urchins, heart urchins, sand dollars, sea biscuits, and pencil urchins. It is divided into two subclasses: '''Perischoechnoidea''', which includes the pencil urchins (or slate-pencil urchins, with very thick, blunt spines), and '''Euechinoidea''', which are the "true" echinoids. The true echinoids, in term, are divided into two groups: ''regular echinoids'' (or regular urchins), which contain the sea urchins, and ''irregular echinoids'' (irregular urchins), which contain the heart urchins, sand dollars, and sea biscuits. All of these have fivefold radial symmetry (pentamerism), but the irregular urchins have secondary bilateral symmetry, with a front and back as well as a top and bottom. Also, while in the regular echinoids the anus is located in the center of the aboral surface (opposite to the mouth, on the dorsal surface), in the irregular echinoids the anus is either posterior or on the oral surface (Follo and Fautin 2001; Freeman 2004).  
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The class '''Echinoidea''' comprises sea urchins, heart urchins, sand dollars, sea biscuits, and pencil urchins. It is divided into two subclasses: '''Perischoechnoidea,''' which includes the pencil urchins (or slate-pencil urchins, with very thick, blunt spines), and '''Euechinoidea,''' which are the "true" echinoids. The true echinoids, in term, are divided into two groups: ''Regular echinoids'' (or regular urchins), which contain the sea urchins, and ''irregular echinoids'' (irregular urchins), which contain the heart urchins, sand dollars, and sea biscuits. All of these have fivefold radial symmetry (pentamerism), but the irregular urchins have secondary bilateral symmetry, with a front and back as well as a top and bottom. Also, while in the regular echinoids the anus is located in the center of the aboral surface (opposite to the mouth, on the dorsal surface), in the irregular echinoids the anus is either posterior or on the oral surface (Follo and Fautin 2001; Freeman 2004).  
  
 
Sea urchins tend to be small, globular, and have their soft internal organs protected by a hard, internal, calcite shell, or test, made of plates (calcitic ossicles) fitting closely together and located under the skin. Their test is round and spiny, typically from 3 to 10 centimeters across. It tends to be arched on the aboral side (dorsal) and flat or concave on the oral side (ventral). Common colors include black and dull shades of green, olive, brown, purple, and red.  
 
Sea urchins tend to be small, globular, and have their soft internal organs protected by a hard, internal, calcite shell, or test, made of plates (calcitic ossicles) fitting closely together and located under the skin. Their test is round and spiny, typically from 3 to 10 centimeters across. It tends to be arched on the aboral side (dorsal) and flat or concave on the oral side (ventral). Common colors include black and dull shades of green, olive, brown, purple, and red.  
  
All sea urchins (indeed all echinoids) are covered by movable spines. These spines are attached to regularly arranged tubercles, which in turn are attached to the test. The [[spine (biology)|spine]]s, which in some species are long and sharp, serve to protect the urchin from [[predator]]s and aid the tube feet in locomotion and digging. The spines can inflict a painful wound on a human who steps on one, but they are not seriously dangerous, and it is not clear that the spines are truly venomous. Typical sea urchins have spines that are 1 to 3 centimeters in length, 1 to 2 millimeters thick, and not terribly sharp. ''[[Diadema antillarum]]'', familiar in the Caribbean, has thin, potentially dangerous spines that can be 10 to 20 centimeters long.
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[[Image:UrchinTestClose.jpg|thumb|left|240px|Closeup of a sea urchin test. In life, a tube foot or gill extends through each of the small holes, and a spine is supported by each of the raised tubercles.]]
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All sea urchins (indeed all echinoids) are covered by movable spines. These spines are attached to regularly arranged tubercles, which in turn are attached to the test. The [[spine (biology)|spine]]s, which in some species are long and sharp, serve to protect the urchin from [[predator]]s and aid the tube feet in locomotion and digging. The spines can inflict a painful wound on a human who steps on one, but they are not seriously dangerous, and it is not clear that the spines are truly venomous. Typical sea urchins have spines that are 1 to 3 centimeters in length, 1 to 2 millimeters thick, and not terribly sharp. ''[[Diadema antillarum]],'' familiar in the Caribbean, has thin, potentially dangerous spines that can be 10 to 20 centimeters long.
  
 
Some sea urchins also have [[pedicellaria]]e between the spines, which are tiny, pincer-like structures that also can be used in defense, or for removal of unwanted parasites, detritus, or microorganisms (Freeman 2004). In some species the pedicellaria are venomous.
 
Some sea urchins also have [[pedicellaria]]e between the spines, which are tiny, pincer-like structures that also can be used in defense, or for removal of unwanted parasites, detritus, or microorganisms (Freeman 2004). In some species the pedicellaria are venomous.
  
The test of sea urchins are pierced by pores through which extend tube feet, which are connected to the water-vascular system. These are known primarily for locomotion, but also are used to trap food, adhere t the substrate, and on the aboral side for respiration and sensation. The tube feet are slender, often sucker-tipped, and extensible.
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The test of sea urchins are pierced by pores through which extend tube feet, which are connected to the water-vascular system. These are known primarily for locomotion, but also are used to trap food, adhere to the substrate, and on the aboral side for [[respiration]] and sensation. The tube feet are slender, often sucker-tipped, and extensible.
  
On the oral surface of the sea urchin, facing the substratum, is a centrally located mouth made up of five united [[calcium carbonate]] teeth or jaws, with a fleshy tongue-like structure within. The entire chewing organ is known as [[Aristotle]]'s lantern, which name comes from Aristotle's accurate description in his ''History of Animals'':
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On the oral surface of the sea urchin, facing the substratum, is a centrally located mouth made up of five united [[calcium carbonate]] teeth or jaws, with a fleshy tongue-like structure within. The entire chewing organ is known as [[Aristotle]]'s lantern, which name comes from Aristotle's accurate description in his ''History of Animals:''
  
:…the urchin has what we mainly call its head and mouth down below, and a place for the issue of the residuum up above. The urchin has, also, five hollow teeth inside, and in the middle of these teeth a fleshy substance serving the office of a tongue. Next to this comes the esophagus, and then the stomach, divided into five parts, and filled with excretion, all the five parts uniting at the anal vent, where the shell is perforated for an outlet... In reality the mouth-apparatus of the urchin is continuous from one end to the other, but to outward appearance it is not so, but looks like a horn lantern with the panes of horn left out. (Tr. [[D'Arcy Thompson]])
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<blockquote>…the urchin has what we mainly call its head and mouth down below, and a place for the issue of the residuum up above. The urchin has, also, five hollow teeth inside, and in the middle of these teeth a fleshy substance serving the office of a tongue. Next to this comes the esophagus, and then the stomach, divided into five parts, and filled with excretion, all the five parts uniting at the anal vent, where the shell is perforated for an outlet… In reality the mouth-apparatus of the urchin is continuous from one end to the other, but to outward appearance it is not so, but looks like a horn lantern with the panes of horn left out (Tr. [[D'Arcy Thompson]]).</blockquote>
  
 
Sea urchins have five pairs of gonads, whereas the irregular echinoids have between two and five (Freeman 2004).
 
Sea urchins have five pairs of gonads, whereas the irregular echinoids have between two and five (Freeman 2004).
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The name ''[[urchin]]'' is an old name for the round spiny [[hedgehog]]s that sea urchins resemble.
 
The name ''[[urchin]]'' is an old name for the round spiny [[hedgehog]]s that sea urchins resemble.
  
==Behavior and diet==
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==Behavior, diet, and ecology==
 
[[Image:SeaDSC01286.JPG|thumb|''[[Echinothrix calamaris]]'', a species of sea urchin. The sphere in the middle of a sea urchin is its anus]]
 
[[Image:SeaDSC01286.JPG|thumb|''[[Echinothrix calamaris]]'', a species of sea urchin. The sphere in the middle of a sea urchin is its anus]]
Sea urchins move slowly, feeding mostly on algae. At first glance, a sea urchin often appears [[sessile]], in other words, incapable of moving. Sometimes the most visible sign of life is the spines, which are attached at their bases to ball-and-socket joints and can be pointed in any direction. In most urchins, a light touch elicits a prompt and visible reaction from the spines, which converge toward the point that has been touched. A sea urchin has no visible eyes, legs, or means of propulsion, but it can move freely over surfaces by means of its adhesive tube feet, working in conjunction with its spines.
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Sea urchins move slowly, with many species feeding mostly on [[algae]]. At first glance, a sea urchin often appears [[sessile]], in other words, incapable of moving. Sometimes the most visible sign of life is the spines, which are attached at their bases to ball-and-socket joints and can be pointed in any direction. In most urchins, a light touch elicits a prompt and visible reaction from the spines, which converge toward the point that has been touched. A sea urchin has no visible eyes, legs, or means of propulsion, but it can move freely over surfaces by means of its adhesive tube feet, working in conjunction with its spines.  
  
Sea urchins feed mainly on [[algae]], but can also feed on a wide range of invertebrates, such as [[mussel]]s, [[sponge]]s, [[brittle star]]s and [[crinoid]]s.  
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Sea urchins feed on [[algae]], sea grasses, seaweeds, and a wide range of [[invertebrate]]s, such as [[mussel]]s, [[sponge]]s, [[brittle star]]s and [[crinoid]]s.  
  
Sea urchin is one of the favorite foods of [[sea otter]]s and are also the main source of nutrition for [[wolf eel]]s. Left unchecked, urchins will devastate their environment, creating what biologists call an [[urchin barren]], devoid of macroalgae and associated fauna. Where sea otters have been re-introduced into [[British Columbia]], the health of the coastal ecosystem has improved dramatically (FOC 2004).
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Sea urchins are eaten by [[crab]]s, [[lobster]]s, [[sea star]]s, [[fish]], [[bird]]s, [[otter]]s, and other mammals (Folo and Fautin 2001). Sea urchin is one of the favorite foods of [[sea otter]]s and are also the main source of nutrition for [[wolf eel]]s. Left unchecked, urchins will devastate their environment, creating what biologists call an [[urchin barren]], devoid of macroalgae and associated fauna. Where sea otters have been re-introduced into [[British Columbia]], the health of the coastal ecosystem has improved dramatically (FOC 2004).
  
 
==Geologic history==
 
==Geologic history==
 
[[Image:Fossil Echinoid Lovenia.jpg|thumb|left|150px|Fossil sea urchin ''Lovenia woodsi'' from the  [[Pliocene]] of [[Australia]].]]
 
[[Image:Fossil Echinoid Lovenia.jpg|thumb|left|150px|Fossil sea urchin ''Lovenia woodsi'' from the  [[Pliocene]] of [[Australia]].]]
The earliest known echinoids are found in the rock of the upper part of the [[Ordovician]] period (''c'' 450 [[million years ago|MYA]]), and they have survived to the present day, where they are a successful and diverse group of organisms. In well-preserved specimens,the spines may be present, but usually only the test is found. Sometimes isolated spines are common as [[fossil]]s. Some echinoids (such as ''Tylocidaris clavigera'', which is found in the [[Cretaceous]] period [[Chalk Formation]] of [[England]]) had very heavy club-shaped spines that would be difficult for an attacking predator to break through and make the echinoid awkward to handle. Such spines are also good for walking on the soft sea-floor.
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The earliest known echinoids are found in the rock of the upper part of the [[Ordovician]] period (''c'' 450 [[million years ago|MYA]]), and they have survived to the present day, where they are a successful and diverse group of organisms. In well-preserved specimens, the spines may be present, but usually only the test is found. Sometimes isolated spines are common as [[fossil]]s. Some echinoids (such as ''Tylocidaris clavigera,'' which is found in the [[Cretaceous]] period [[Chalk Formation]] of [[England]]) had very heavy club-shaped spines that would be difficult for an attacking predator to break through and make the echinoid awkward to handle. Such spines are also good for walking on the soft sea-floor.
  
 
[[Image:Echinoids-castleHainNC.jpg|thumb|right|150px|[[Cretaceous]] echinoids from Castle Hayne quarry, [[North Carolina]], USA]]
 
[[Image:Echinoids-castleHainNC.jpg|thumb|right|150px|[[Cretaceous]] echinoids from Castle Hayne quarry, [[North Carolina]], USA]]
Complete fossil echinoids from the [[Paleozoic]] era are generally rare, usually consisting of isolated spines and small clusters of scattered plates from crushed individuals. Most specimens occur in rocks from the [[Devonian]] and [[Carboniferous]] periods. The shallow water limestones from the Ordovician and [[Silurian]] periods of [[Estonia]] are famous for the echinoids found there. The Paleozoic echinoids probably inhabited relatively quiet waters. Because of their thin test, they would certainly not have survived in the turbulent wave-battered coastal waters inhabited by many modern echinoids today. During the upper part of the Carboniferous period, there was a marked decline in echinoid diversity, and this trend continued into the Permian period. They neared extinction at the end of the Paleozoic era, with just six species known from the [[Permian]] period. Only two separate lineages survived the massive extinction of this period and into the Triassic: the genus ''Miocidaris'', which gave rise to the modern cidaroids (pencil urchins), and the ancestor that gave rise to the euechinoids. By the upper part of the [[Triassic]] period, their numbers began to increase again.  
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Complete fossil echinoids from the [[Paleozoic]] era are generally rare, usually consisting of isolated spines and small clusters of scattered plates from crushed individuals. Most specimens occur in rocks from the [[Devonian]] and [[Carboniferous]] periods. The shallow water limestones from the Ordovician and [[Silurian]] periods of [[Estonia]] are famous for the echinoids found there. The Paleozoic echinoids probably inhabited relatively quiet waters. Because of their thin test, they would certainly not have survived in the turbulent wave-battered coastal waters inhabited by many modern echinoids today. During the upper part of the Carboniferous period, there was a marked decline in echinoid diversity, and this trend continued into the Permian period. They neared extinction at the end of the Paleozoic era, with just six species known from the [[Permian]] period. Only two separate lineages survived the massive extinction of this period and into the Triassic: The genus ''Miocidaris,'' which gave rise to the modern cidaroids (pencil urchins), and the ancestor that gave rise to the euechinoids. By the upper part of the [[Triassic]] period, their numbers began to increase again.  
  
The cidaroids have changed very little since their modern design was established in the Late Triassic and are today considered more or less as living fossils. [[Image:Saddle Wrasse - Thalassoma duperrey feeding on sea urchin.jpg|250px|thumb|Saddle Wrasse, ''Thalassoma duperrey'' is feeding on sea urchin]]
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The cidaroids have changed very little since their modern design was established in the Late Triassic and are today considered more or less as living fossils. [[Image:Saddle Wrasse - Thalassoma duperrey feeding on sea urchin.jpg|250px|thumb|Saddle Wrasse, ''Thalassoma duperrey,'' is feeding on sea urchin]]
The euechinoids, on the other hand, diversified into new lineages throughout the [[Jurassic]] period and into the Cretaceous period. From them emerged the first irregular echinoids (superorder Atelostomata) during the early Jurassic, and when including the other superorder (Gnathostomata) or irregular urchins which evolved independently later, they now represent 47% of all present species of echinoids thanks to their adaptive breakthroughs in both habit and feeding strategy, which allowed them to exploit habitats and food sources unavailable to regular echinoids.  
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The euechinoids, on the other hand, diversified into new lineages throughout the [[Jurassic]] period and into the Cretaceous period. From them emerged the first irregular echinoids (superorder Atelostomata) during the early Jurassic, and when including the other superorder (Gnathostomata) or irregular urchins which evolved independently later, they now represent 47 percent of all present species of echinoids thanks to their adaptive breakthroughs in both habit and feeding strategy, which allowed them to exploit habitats and food sources unavailable to regular echinoids.  
  
During the [[Mesozoic]] and [[Cenozoic]] eras, the echinoids flourished. While most echinoid fossils are restricted to certain localities and formations, where they do occur, they are quite often abundant. An example of this is ''Enallaster'', which may be collected by the thousands in certain outcrops of [[limestone]] from the Cretaceous period in [[Texas]]. Many fossils of the [[Late Jurassic]] ''[[Plesiocidaris]]'' still have the spines attached.
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During the [[Mesozoic]] and [[Cenozoic]] eras, the echinoids flourished. While most echinoid fossils are restricted to certain localities and formations, where they do occur, they are quite often abundant. An example of this is ''Enallaster,'' which may be collected by the thousands in certain outcrops of [[limestone]] from the Cretaceous period in [[Texas]]. Many fossils of the [[Late Jurassic]] ''[[Plesiocidaris]]'' still have the spines attached.
  
 
Some echinoids, such as ''Micraster'' which is found in the Cretaceous period [[Chalk Formation]] of [[England]] and [[France]], serve as zone or [[Index fossil|index]] fossils. Because they evolved rapidly over time, such fossils are useful in enabling geologists to date the rocks in which they are found. However, most echinoids are not abundant enough and may be too limited in their geographic distribution to serve as zone fossils.
 
Some echinoids, such as ''Micraster'' which is found in the Cretaceous period [[Chalk Formation]] of [[England]] and [[France]], serve as zone or [[Index fossil|index]] fossils. Because they evolved rapidly over time, such fossils are useful in enabling geologists to date the rocks in which they are found. However, most echinoids are not abundant enough and may be too limited in their geographic distribution to serve as zone fossils.
  
In the early [[Tertiary]] (''c'' 65 to 1.8 MYA), sand dollars (order Clypeasteroida) arose. Their distinctive flattened test and tiny spines were adapted to life on or under loose sand. They form the newest branch on the echinoid tree.
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In the early [[Tertiary]] (c. 65 to 1.8 MYA), sand dollars (order Clypeasteroida) arose. Their distinctive flattened test and tiny spines were adapted to life on or under loose sand. They form the newest branch on the echinoid tree.
  
 
==Model organism==
 
==Model organism==
 
 
Sea urchins are one of the traditional [[model organisms]] in [[developmental biology]]. The use of sea urchins in this context originates from the 1800s, when the embryonic development of the sea urchins was noticed to be particularly easily viewed by microscopy. Sea urchins were the first species in which the [[sperm]] cells were proven to play an important role in [[reproduction]] by fertilizing the [[ovum]].
 
Sea urchins are one of the traditional [[model organisms]] in [[developmental biology]]. The use of sea urchins in this context originates from the 1800s, when the embryonic development of the sea urchins was noticed to be particularly easily viewed by microscopy. Sea urchins were the first species in which the [[sperm]] cells were proven to play an important role in [[reproduction]] by fertilizing the [[ovum]].
  
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Image:urchin003.jpg|Group of black, long-spined Caribbean sea urchins, ''[[Diadema antillarum]]'' (Philippi)
 
Image:urchin003.jpg|Group of black, long-spined Caribbean sea urchins, ''[[Diadema antillarum]]'' (Philippi)
 
Image:Sea urchin eggs.jpg|Sea urchin [[roe]].
 
Image:Sea urchin eggs.jpg|Sea urchin [[roe]].
Image:Zeeegel3.jpg|Sea urchin test. Each white band is the location of a row of tube feet; each pair of white bands is called an ambulacrum. There are five such ambulacra; the fivefold symmetry reveals a kinship with starfish.
 
Image:UrchinTestClose.jpg|Closeup of a sea urchin test. In life, a tube foot or gill extends through each of the small holes, and a spine is supported by each of the raised tubercles.
 
 
Image:Seeigel-Saugfuesse(Galicien2005).jpg|Sea urchins have adhesive tube feet.
 
Image:Seeigel-Saugfuesse(Galicien2005).jpg|Sea urchins have adhesive tube feet.
 
Image:Sea Urchin off Florida coast 2007-08-06.JPG|Sea urchin in a reef off of the Florida coast.
 
Image:Sea Urchin off Florida coast 2007-08-06.JPG|Sea urchin in a reef off of the Florida coast.
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</gallery>
 
</gallery>
  
==References and further reading==
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==References==
* {{aut|Smith, Andrew B.}} (1984), ''Echinoid Palaeobiology (Special topics in palaeontology)''. London: Allen & Unwin. ISBN 0-04-563001-1
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* Fisheries and Oceans Canada (FOC). 2004. [http://www.dfo-mpo.gc.ca/species-especes/species/species_seaOtter_e.asp Sea otter.] ''Fisheries and Oceans Canada''. Retrieved October 12, 2008.
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* Follo, J., and D. Fautin. 2001. [http://animaldiversity.ummz.umich.edu/site/accounts/information/Echinoidea.html Class Echinoidea: Heart urchins, sand dollars, and sea urchins.] ''Animal Diversity Web'' (online). Retrieved October 12, 2008.
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* Freeman, S. M. 2004. Echinoidea. In B. Grzimek, S. F. Craig, D. A. Thoney, N. Schlager, and M. Hutchins. ''Grzimek's Animal Life Encyclopedia,'' 2nd edition. Detroit, MI: Thomson/Gale. ISBN 0787657786.
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* Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2008. [http://animaldiversity.ummz.umich.edu/site/accounts/classification/Echinoidea.html#Echinoidea Class Echinoidea (heart urchins, sand dollars, and sea urchins).] ''Animal Diversity Web'' (online). Retrieved October 12, 2008.
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* Rast, J. P. et al. 2006. Genomic insights into the immune system of the sea urchin. ''Science'' 314(5801): 952-6.
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* Smith, A. B., 1984. ''Echinoid Palaeobiology (Special topics in palaeontology)''. London: Allen & Unwin. ISBN 0045630011.
  
http://animaldiversity.ummz.umich.edu/site/accounts/classification/Echinoidea.html#Echinoidea Animal Diversity Web Classification of the Echinoidea]
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==External links==
Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2008. The Animal Diversity Web (online). Accessed October 12, 2008 at http://animaldiversity.org. 
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All links retrieved January 25, 2023.  
Class Echinoidea
 
(heart urchins, sand dollars, and sea urchins)
 
  
http://animaldiversity.ummz.umich.edu/site/accounts/information/Echinoidea.html
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*[http://news.bbc.co.uk/2/hi/science/nature/6130244.stm Spiny creature's genome insight]
Class Echinoidea: heart urchins, sand dollars, and sea urchins
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*[http://www.nhm.ac.uk/research-curation/projects/echinoid-directory/morphology/JPEG/LANTERN.jpg lantern.jpg] A labeled diagram of the sea urchin's Aristotle's lantern.
Follo, J. and D. Fautin. 2001. "Echinoidea" (On-line), Animal Diversity Web. Accessed October 12, 2008 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Echinoidea.html
 
 
 
 
 
 
 
http://whale.wheelock.edu/archives/ask99/0388.html#The Ocean Alliance giving advice on sea urchin cleaning]
 
 
 
 
 
Bob Goemans http://www.saltcorner.com/sections/zoo/inverts/echinoderms/urchins/urchins.htm Urchins
 
Class Echinoidea  Animal Library
 
  
Fisheries and Oceans Canada. 2004. .<ref name=dfo>{{cite web|url=http://www.dfo-mpo.gc.ca/species-especes/species/species_seaOtter_e.asp|title=Aquatic Species at Risk - Species Profile - Sea Otter|publisher=[[Fisheries and Oceans Canada]]|accessdate=2007-11-29}}</ref>
 
  
<ref>Rast, JP et al. ''Genomic insights into the immune system of the sea urchin.'' Science. 2006 Nov 10;314(5801):952-6.</ref>.
 
 
==External links==
 
{{commons|Sea urchin}}
 
*[http://seaurchin.org/ Sea Urchin Harvesters Association - California] Also, (604) 524-0322.
 
*[http://www.nhm.ac.uk/research-curation/projects/echinoid-directory The Echinoid Directory] from the [[Natural History Museum]].
 
*[http://nlbif.eti.uva.nl/bis/echinodermata.php?menuentry=zoeken&selected=boom&echinodermata-taxon=class&echinodermata-node=Echinoidea&echinodermata-treepath=none,Animalia,Echinodermata,Echinoidea&collapsed=false#Echinoidea Echinoids of the North Sea]
 
*[http://www.livescience.com/animalworld/061109_urchin_relatives.html 70% of Sea Urchin Genes Have a Human Counterpart] — Sequencing confirms that sea urchins are more closely related to humans than fruit flies (LiveScience.com, November 2006).
 
*[http://news.bbc.co.uk/2/hi/science/nature/6130244.stm Spiny creature's genome insight]
 
*[http://www.thesea.org/TheSea/purple_sea_urchin.html 80 gallon tank video] of purple sea urchin
 
*[http://www.echinoids.nl/ Echinoids.nl]
 
*[http://www.nhm.ac.uk/research-curation/projects/echinoid-directory/morphology/JPEG/LANTERN.jpg lantern.jpg] A labeled diagram of the sea urchin's Aristotle's lantern.
 
*[http://askabiologist.asu.edu/research/urchins/aristotle.html aristotle.htm] Who is this person Aristotle and what about this lantern?
 
*[http://www.emilydamstra.com/portfolio2.php?illid=56 www.emilydamstra.com] Illustration of the musculature of an Aristotle's lantern.
 
*[http://virtualurchin.stanford.edu/urchinanatomy.swf Urchin Anatomy] a flash about the anatomy of the sea urchin
 
  
 
[[Category:Life sciences]]
 
[[Category:Life sciences]]

Latest revision as of 17:33, 25 January 2023

Sea urchin
Sea urchins, Sterechinus neumayeri
Sea urchins, Sterechinus neumayeri
Scientific classification
Kingdom: Animalia
Phylum: Echinodermata
Subphylum: Echinozoa
Class: Echinoidea
Leske, 1778
Subclasses
  • Subclass Perischoechinoidea
    • Order Cidaroida (pencil urchins)
  • Subclass Euechinoidea
    • Superorder Atelostomata
      • Order Cassiduloida
      • Order Spatangoida (heart urchins)
    • Superorder Diadematacea
      • Order Diadematoida
      • Order Echinothurioida
      • Order Pedinoida
    • Superorder Echinacea
      • Order Arbacioida
      • Order Echinoida
      • Order Phymosomatoida
      • Order Salenioida
      • Order Temnopleuroida
    • Superorder Gnathostomata
      • Order Clypeasteroida (sand dollars)
      • Order Holectypoida

Sea urchin is the common name for various spiky echinoderms within the class Echinoidea, characterized by pentamerous radial symmetry; a hard calcareous shell, or test, made of regularly arranged, tightly fused plates through which rows of slender, extensible tube feet extend; movable spines covering the shell; and a mouth on the lower side, facing the substratum, with a complex chewing structure composed of five jaws (Aristotle's lantern).

In a more general sense of the term, sea urchin sometimes is used as the common name for all members of Echinoidea. However, more specifically, it is used to refer to the "regular echinoids" within subclass Euechinoidea that are pentaradially symmetrical and have their anus located on the aboral surface (opposite to the mouth). The term sea urchin, thus, does not normally include the pencil urchins in subclass Perishoechinoidea, nor the "irregular echinoids" within Euechinoidea, which include the heart urchins and sand dollars. The later tend to be characterized by a secondary bilateral symmetry and a more flattened and oval test, with very short spines, and with the anus either posterior or on the oral surface.

Sea urchins are found in oceans all over the world. They play an important role in marine food chains, consuming algae and various invertebrates, and being consumed by crabs, sea stars, fishes, mammals, and birds. When predators such as sea otters disappear from the ecosystem, the unchecked sea urchins can devastate their environment, making an "urchin barren."

For humans, sea urchins are harvested and served as a delicacy. Their roe and gonads are frequently consumed raw, or briefly cooked, with species such as the red sea urchin (Strongylocentrotus francis-canus), green urchin (S. droebachiensis), and purple sea urchin (S. purpuratus) among those favored for their roe (Freeman 2004). They are a frequent model organism for developmental and immunological studies.

Overview and description

Sea urchin test. Each white band is the location of a row of tube feet; each pair of white bands is called an ambulacrum. There are five such ambulacra; the fivefold symmetry reveals a kinship with starfish.

Sea urchins are members of the phylum Echinodermata, which also includes starfish, sea cucumbers, brittle stars, and crinoids. Like other echinoderms they have fivefold symmetry (called pentamerism) and move by means of hundreds of tiny, transparent, adhesive "tube feet." The pentamerous symmetry is not obvious at a casual glance but is easily seen in the dried shell or test of the urchin.

The class Echinoidea comprises sea urchins, heart urchins, sand dollars, sea biscuits, and pencil urchins. It is divided into two subclasses: Perischoechnoidea, which includes the pencil urchins (or slate-pencil urchins, with very thick, blunt spines), and Euechinoidea, which are the "true" echinoids. The true echinoids, in term, are divided into two groups: Regular echinoids (or regular urchins), which contain the sea urchins, and irregular echinoids (irregular urchins), which contain the heart urchins, sand dollars, and sea biscuits. All of these have fivefold radial symmetry (pentamerism), but the irregular urchins have secondary bilateral symmetry, with a front and back as well as a top and bottom. Also, while in the regular echinoids the anus is located in the center of the aboral surface (opposite to the mouth, on the dorsal surface), in the irregular echinoids the anus is either posterior or on the oral surface (Follo and Fautin 2001; Freeman 2004).

Sea urchins tend to be small, globular, and have their soft internal organs protected by a hard, internal, calcite shell, or test, made of plates (calcitic ossicles) fitting closely together and located under the skin. Their test is round and spiny, typically from 3 to 10 centimeters across. It tends to be arched on the aboral side (dorsal) and flat or concave on the oral side (ventral). Common colors include black and dull shades of green, olive, brown, purple, and red.

Closeup of a sea urchin test. In life, a tube foot or gill extends through each of the small holes, and a spine is supported by each of the raised tubercles.

All sea urchins (indeed all echinoids) are covered by movable spines. These spines are attached to regularly arranged tubercles, which in turn are attached to the test. The spines, which in some species are long and sharp, serve to protect the urchin from predators and aid the tube feet in locomotion and digging. The spines can inflict a painful wound on a human who steps on one, but they are not seriously dangerous, and it is not clear that the spines are truly venomous. Typical sea urchins have spines that are 1 to 3 centimeters in length, 1 to 2 millimeters thick, and not terribly sharp. Diadema antillarum, familiar in the Caribbean, has thin, potentially dangerous spines that can be 10 to 20 centimeters long.

Some sea urchins also have pedicellariae between the spines, which are tiny, pincer-like structures that also can be used in defense, or for removal of unwanted parasites, detritus, or microorganisms (Freeman 2004). In some species the pedicellaria are venomous.

The test of sea urchins are pierced by pores through which extend tube feet, which are connected to the water-vascular system. These are known primarily for locomotion, but also are used to trap food, adhere to the substrate, and on the aboral side for respiration and sensation. The tube feet are slender, often sucker-tipped, and extensible.

On the oral surface of the sea urchin, facing the substratum, is a centrally located mouth made up of five united calcium carbonate teeth or jaws, with a fleshy tongue-like structure within. The entire chewing organ is known as Aristotle's lantern, which name comes from Aristotle's accurate description in his History of Animals:

…the urchin has what we mainly call its head and mouth down below, and a place for the issue of the residuum up above. The urchin has, also, five hollow teeth inside, and in the middle of these teeth a fleshy substance serving the office of a tongue. Next to this comes the esophagus, and then the stomach, divided into five parts, and filled with excretion, all the five parts uniting at the anal vent, where the shell is perforated for an outlet… In reality the mouth-apparatus of the urchin is continuous from one end to the other, but to outward appearance it is not so, but looks like a horn lantern with the panes of horn left out (Tr. D'Arcy Thompson).

Sea urchins have five pairs of gonads, whereas the irregular echinoids have between two and five (Freeman 2004).

The name urchin is an old name for the round spiny hedgehogs that sea urchins resemble.

Behavior, diet, and ecology

Echinothrix calamaris, a species of sea urchin. The sphere in the middle of a sea urchin is its anus

Sea urchins move slowly, with many species feeding mostly on algae. At first glance, a sea urchin often appears sessile, in other words, incapable of moving. Sometimes the most visible sign of life is the spines, which are attached at their bases to ball-and-socket joints and can be pointed in any direction. In most urchins, a light touch elicits a prompt and visible reaction from the spines, which converge toward the point that has been touched. A sea urchin has no visible eyes, legs, or means of propulsion, but it can move freely over surfaces by means of its adhesive tube feet, working in conjunction with its spines.

Sea urchins feed on algae, sea grasses, seaweeds, and a wide range of invertebrates, such as mussels, sponges, brittle stars and crinoids.

Sea urchins are eaten by crabs, lobsters, sea stars, fish, birds, otters, and other mammals (Folo and Fautin 2001). Sea urchin is one of the favorite foods of sea otters and are also the main source of nutrition for wolf eels. Left unchecked, urchins will devastate their environment, creating what biologists call an urchin barren, devoid of macroalgae and associated fauna. Where sea otters have been re-introduced into British Columbia, the health of the coastal ecosystem has improved dramatically (FOC 2004).

Geologic history

Fossil sea urchin Lovenia woodsi from the Pliocene of Australia.

The earliest known echinoids are found in the rock of the upper part of the Ordovician period (c 450 MYA), and they have survived to the present day, where they are a successful and diverse group of organisms. In well-preserved specimens, the spines may be present, but usually only the test is found. Sometimes isolated spines are common as fossils. Some echinoids (such as Tylocidaris clavigera, which is found in the Cretaceous period Chalk Formation of England) had very heavy club-shaped spines that would be difficult for an attacking predator to break through and make the echinoid awkward to handle. Such spines are also good for walking on the soft sea-floor.

Cretaceous echinoids from Castle Hayne quarry, North Carolina, USA

Complete fossil echinoids from the Paleozoic era are generally rare, usually consisting of isolated spines and small clusters of scattered plates from crushed individuals. Most specimens occur in rocks from the Devonian and Carboniferous periods. The shallow water limestones from the Ordovician and Silurian periods of Estonia are famous for the echinoids found there. The Paleozoic echinoids probably inhabited relatively quiet waters. Because of their thin test, they would certainly not have survived in the turbulent wave-battered coastal waters inhabited by many modern echinoids today. During the upper part of the Carboniferous period, there was a marked decline in echinoid diversity, and this trend continued into the Permian period. They neared extinction at the end of the Paleozoic era, with just six species known from the Permian period. Only two separate lineages survived the massive extinction of this period and into the Triassic: The genus Miocidaris, which gave rise to the modern cidaroids (pencil urchins), and the ancestor that gave rise to the euechinoids. By the upper part of the Triassic period, their numbers began to increase again.

The cidaroids have changed very little since their modern design was established in the Late Triassic and are today considered more or less as living fossils.

Saddle Wrasse, Thalassoma duperrey, is feeding on sea urchin

The euechinoids, on the other hand, diversified into new lineages throughout the Jurassic period and into the Cretaceous period. From them emerged the first irregular echinoids (superorder Atelostomata) during the early Jurassic, and when including the other superorder (Gnathostomata) or irregular urchins which evolved independently later, they now represent 47 percent of all present species of echinoids thanks to their adaptive breakthroughs in both habit and feeding strategy, which allowed them to exploit habitats and food sources unavailable to regular echinoids.

During the Mesozoic and Cenozoic eras, the echinoids flourished. While most echinoid fossils are restricted to certain localities and formations, where they do occur, they are quite often abundant. An example of this is Enallaster, which may be collected by the thousands in certain outcrops of limestone from the Cretaceous period in Texas. Many fossils of the Late Jurassic Plesiocidaris still have the spines attached.

Some echinoids, such as Micraster which is found in the Cretaceous period Chalk Formation of England and France, serve as zone or index fossils. Because they evolved rapidly over time, such fossils are useful in enabling geologists to date the rocks in which they are found. However, most echinoids are not abundant enough and may be too limited in their geographic distribution to serve as zone fossils.

In the early Tertiary (c. 65 to 1.8 MYA), sand dollars (order Clypeasteroida) arose. Their distinctive flattened test and tiny spines were adapted to life on or under loose sand. They form the newest branch on the echinoid tree.

Model organism

Sea urchins are one of the traditional model organisms in developmental biology. The use of sea urchins in this context originates from the 1800s, when the embryonic development of the sea urchins was noticed to be particularly easily viewed by microscopy. Sea urchins were the first species in which the sperm cells were proven to play an important role in reproduction by fertilizing the ovum.

With the recent sequencing of the sea urchin genome, homology has been found between sea urchin and vertebrate immune system-related genes. Sea urchins code for at least 222 Toll-like receptor (TLR) genes and over 200 genes related to the Nod-like-receptor (NLR) family found in vertebrates (Rast et al. 2006). This has made the sea urchin a valuable model organism for immunologists to study the development of innate immunity.

Gallery

References
ISBN links support NWE through referral fees

  • Fisheries and Oceans Canada (FOC). 2004. Sea otter. Fisheries and Oceans Canada. Retrieved October 12, 2008.
  • Follo, J., and D. Fautin. 2001. Class Echinoidea: Heart urchins, sand dollars, and sea urchins. Animal Diversity Web (online). Retrieved October 12, 2008.
  • Freeman, S. M. 2004. Echinoidea. In B. Grzimek, S. F. Craig, D. A. Thoney, N. Schlager, and M. Hutchins. Grzimek's Animal Life Encyclopedia, 2nd edition. Detroit, MI: Thomson/Gale. ISBN 0787657786.
  • Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2008. Class Echinoidea (heart urchins, sand dollars, and sea urchins). Animal Diversity Web (online). Retrieved October 12, 2008.
  • Rast, J. P. et al. 2006. Genomic insights into the immune system of the sea urchin. Science 314(5801): 952-6.
  • Smith, A. B., 1984. Echinoid Palaeobiology (Special topics in palaeontology). London: Allen & Unwin. ISBN 0045630011.

External links

All links retrieved January 25, 2023.

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