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An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include
starfish Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
,
brittle star Brittle stars, serpent stars, or ophiuroids (; ; referring to the serpent-like arms of the brittle star) are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomot ...
s, sea urchins,
sand dollar Sand dollars (also known as a sea cookie or snapper biscuit in New Zealand, or pansy shell in South Africa) are species of flat, burrowing sea urchins belonging to the order Clypeasteroida. Some species within the order, not quite as flat, are ...
s, and sea cucumbers, as well as the sea lilies or "stone lilies". Adult echinoderms are found on the sea bed at every ocean depth, from the
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species of ...
to the
abyssal zone The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. "Abyss" derives from the Greek word , meaning bottomless. At depths of , this zone remains in perpetual darkness. It covers 83% of the total area of the ocean an ...
. The phylum contains about 7,000 living
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
, making it the second-largest grouping of deuterostomes, after the
chordates A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...
. Echinoderms are the largest entirely marine phylum. The first definitive echinoderms appeared near the start of the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
. The echinoderms are important both ecologically and geologically. Ecologically, there are few other groupings so abundant in the biotic desert of the deep sea, as well as shallower oceans. Most echinoderms are able to reproduce asexually and regenerate tissue, organs, and limbs; in some cases, they can undergo complete regeneration from a single limb. Geologically, the value of echinoderms is in their
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in th ...
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s, which are major contributors to many limestone formations, and can provide valuable clues as to the geological environment. They were the most used species in regenerative research in the 19th and 20th centuries. Further, some scientists hold that the radiation of echinoderms was responsible for the Mesozoic Marine Revolution.


Taxonomy and evolution

The name echinoderm is . Echinoderms are
bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
ns, meaning that their ancestors were mirror-symmetric. Among the bilaterians, they belong to the deuterostome division, meaning that the
blastopore Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals the blastocyst is reorganized into a multilayered structure known as the gastrula. Be ...
, the first opening to form during embryo development, becomes the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, de ...
instead of the mouth. The characteristics of adult echinoderms are the possession of a
water vascular system The water vascular system is a hydraulic system used by echinoderms, such as sea stars and sea urchins, for locomotion, food and waste transportation, and respiration. The system is composed of canals connecting numerous tube feet. Echinoderms mov ...
with external
tube feet Tube feet (technically podia) are small active tubular projections on the oral face of an echinoderm, whether the arms of a starfish, or the undersides of sea urchins, sand dollars and sea cucumbers; they are more discreet though present on brit ...
and a
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
endoskeleton consisting of
ossicles The ossicles (also called auditory ossicles) are three bones in either middle ear that are among the smallest bones in the human body. They serve to transmit sounds from the air to the fluid-filled labyrinth ( cochlea). The absence of the audit ...
connected by a mesh of collagen fibres.


Phylogeny

Historically, taxonomists believed that the Ophiuroidea were sister to the Asteroidea, or that they were sister to the (Holothuroidea + Echinoidea). However, a 2014 analysis of 219 genes from all classes of echinoderms revised the phylogenetic tree. An independent analysis in 2015 of RNA transcriptomes from 23 species across all classes of echinoderms gave the same tree. ; External phylogeny The context of the echinoderms within the Bilateria is: ; Internal phylogeny


Diversity

There are about 7,000
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extin ...
species of echinoderm as well as about 13,000 extinct species. All echinoderms are marine, but they are found in habitats ranging from shallow intertidal areas to abyssal depths. Two main subdivisions are traditionally recognised: the more familiar
motile Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
Eleutherozoa Eleutherozoa is a proposed subphylum of echinoderms. They are mobile animals with the mouth directed towards the substrate. They usually have a madreporite, tube feet, and moveable spines of some sort, and some have Tiedemann's bodies on the ...
, which encompasses the Asteroidea (
starfish Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
, with some 1,745 species), Ophiuroidea (
brittle stars Brittle stars, serpent stars, or ophiuroids (; ; referring to the serpent-like arms of the brittle star) are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomot ...
, with around 2,300 species), Echinoidea ( sea urchins and
sand dollar Sand dollars (also known as a sea cookie or snapper biscuit in New Zealand, or pansy shell in South Africa) are species of flat, burrowing sea urchins belonging to the order Clypeasteroida. Some species within the order, not quite as flat, are ...
s, with some 900 species) and Holothuroidea ( sea cucumbers, with about 1,430 species); and the Pelmatozoa, some of which are
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
while others are motile. These consist of the Crinoidea ( feather stars and sea lilies, with around 580 species) and the extinct
blastoids Blastoids (class Blastoidea) are an extinct type of stemmed echinoderm, often referred to as sea buds. They first appear, along with many other echinoderm classes, in the Ordovician period, and reached their greatest diversity in the Mississi ...
and
Paracrinoid Paracrinoidea is an extinct class of blastozoan echinoderms. They lived in shallow seas during the Early Ordovician through the Early Silurian. While blastozoans are usually characterized by types of respiratory structures present, it is not cle ...
s. File:Ophionereis reticulata 1.jpg, A brittle star, ''
Ophionereis reticulata ''Ophionereis reticulata'', the reticulated brittle star, is a brittle star in the family Ophionereididae. It is found in shallow parts of the western Atlantic, Caribbean Sea and Gulf of Mexico. Description Like other brittle stars, ''Ophionerei ...
'' File:Sea cucumber at Pulau Redang.jpg, A sea cucumber from Malaysia File:Nerr0878.jpg, Starfish of varied colours File:Strongylocentrotus purpuratus 1.jpg, A sea urchin, ''
Strongylocentrotus purpuratus ''Strongylocentrotus purpuratus'', the purple sea urchin, lives along the eastern edge of the Pacific Ocean extending from Ensenada, Mexico, to British Columbia, Canada. This sea urchin species is deep purple in color, and lives in lower int ...
'' File:Crinoid on the reef of Batu Moncho Island.JPG,
Crinoid Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which ar ...
on a coral reef


Fossil history

The oldest candidate echinoderm fossil is ''
Arkarua ''Arkarua adami'' is a small, Precambrian disk-like fossil with a raised center, a number of radial ridges on the rim, and a five-pointed central depression marked with radial lines of five small dots from the middle of the disk center. Fossils ...
'' from the Precambrian of Australia. These fossils are disc-like, with radial ridges on the rim and a five-pointed central depression marked with radial lines. However, the fossils have no
stereom Stereom is a calcium carbonate material that makes up the internal skeletons found in all echinoderms, both living and fossilized forms. It is a sponge-like porous structure which, in a sea urchin may be 50% by volume living cells, and the rest b ...
or internal structure indicating a water vascular system, so they cannot be conclusively identified. The first universally accepted echinoderms appear in the
Lower Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
period; asterozoans appeared in the
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...
, while the crinoids were a dominant group in the Paleozoic. Echinoderms left behind an extensive fossil record. It is hypothesised that the ancestor of all echinoderms was a simple, motile, bilaterally symmetrical animal with a mouth, gut and anus. This ancestral organism adopted an attached mode of life with suspension feeding, and developed radial symmetry. Even so, the larvae of all echinoderms are bilaterally symmetrical, and all develop radial symmetry at metamorphosis. Like their ancestor, the starfish and crinoids still attach themselves to the seabed while changing to their adult form. The first echinoderms were non-motile, but evolved into animals able to move freely. These soon developed endoskeletal plates with stereom structure, and external ciliary grooves for feeding. The Paleozoic echinoderms were globular, attached to the substrate and were orientated with their oral surfaces facing upwards. These early echinoderms had
ambulacral groove Ambulacral is a term typically used in the context of anatomical parts of the phylum Echinodermata or class Asteroidea and Edrioasteroidea. Echinoderms can have ambulacral parts that include ossicles, plates, spines, and suckers. For example, se ...
s extending down the side of the body, fringed on either side by brachioles, like the pinnules of a modern crinoid. Eventually, except for the crinoids, all the classes of echinoderms reversed their orientation to become mouth-downward. Before this happened, the podia probably had a feeding function, as they do in the crinoids today. The locomotor function of the podia came later, when the re-orientation of the mouth brought the podia into contact with the substrate for the first time. :Further information: '' Dibrachicystis'' File:EarlyEchinoderms NT.jpg, Early echinoderms '' Ctenoimbricata'', '' Ctenocystis'', '' Gogia'', '' Protocintus'' and '' Rhenocystis'' File:Echinosphaerites.JPG, The Ordovician cystoid ''
Echinosphaerites ''Echinosphaerites'' is a genus of rhombiferan cystoid echinoderms that lived in the Early to Middle Ordovician of North America and Europe (Bockelie, 1981). Biology Echinosphaerites had branched biserial brachioles which is rare for species ...
'' from northeastern Estonia File:fossile-seelilie.jpg, Fossil
crinoid Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which ar ...
crowns File:Hyperoblastus.jpg, Calyx of ''Hyperoblastus'', a
blastoid Blastoids (class Blastoidea) are an extinct type of stemmed echinoderm, often referred to as sea buds. They first appear, along with many other echinoderm classes, in the Ordovician period, and reached their greatest diversity in the Mississi ...
from the Devonian of Wisconsin


Anatomy and physiology

Echinoderms evolved from animals with
bilateral symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pl ...
. Although adult echinoderms possess pentaradial symmetry, their larvae are ciliated, free-swimming organisms with bilateral symmetry. Later, during metamorphosis, the left side of the body grows at the expense of the right side, which is eventually absorbed. The left side then grows in a pentaradially symmetric fashion, in which the body is arranged in five parts around a central axis. Within the
Asterozoa The Asterozoa are a subphylum in the phylum Echinodermata. Characteristics include a star-shaped body and radially divergent axes of symmetry. The subphylum includes the class Asteroidea (the starfish), the class Ophiuroidea (the brittle stars ...
, there can be a few exceptions from the rule. Most starfish in the genus '' Leptasterias'' have six arms, although five-armed individuals can occur. The
Brisingida The Brisingids are deep-sea-dwelling starfish in the order Brisingida. Description These starfish have between 6 to 18 long, attenuated arms which they use for suspension feeding. Other characteristics include a single series of marginals, a f ...
too contain some six-armed species. Amongst the brittle stars, six-armed species such as ''Ophiothela danae'', ''
Ophiactis savignyi ''Ophiactis savignyi'' is a species of brittle star in the family Ophiactidae, commonly known as Savigny's brittle star or the little brittle star. It occurs in the tropical and subtropical parts of all the world's oceans and is thought to be t ...
'', and ''Ophionotus hexactis'' exists, and ''Ophiacantha vivipara'' often has more than six. Echinoderms have secondary radial symmetry in portions of their body at some stage of life, most likely an adaptation to a sessile or slow-moving existence. Many crinoids and some seastars are symmetrical in multiples of the basic five; starfish such as ''
Labidiaster annulatus ''Labidiaster annulatus'', the Antarctic sun starfish or wolftrap starfish is a species of starfish in the family Heliasteridae. It is found in the cold waters around Antarctica and has a large number of slender, flexible rays. Description ' ...
'' possess up to fifty arms, while the sea-lily '' Comaster schlegelii'' has two hundred.


Skin and skeleton

Echinoderms have a mesodermal skeleton in the dermis, composed of
calcite Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratc ...
-based plates known as
ossicles The ossicles (also called auditory ossicles) are three bones in either middle ear that are among the smallest bones in the human body. They serve to transmit sounds from the air to the fluid-filled labyrinth ( cochlea). The absence of the audit ...
. If solid, these would form a heavy skeleton, so they have a sponge-like porous structure known as stereom. Ossicles may be fused together, as in the test of sea urchins, or may
articulate Articulate may refer to: * '' Articulate!'', a board game in which players describe words from different categories * Articulate brachiopods, brachiopods with toothed hinges and simple opening and closing muscles * Articulate sound, to move the ...
to form flexible joints as in the arms of sea stars, brittle stars and crinoids. The ossicles may bear external projections in the form of spines, granules or warts and they are supported by a tough
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
. Skeletal elements are sometimes deployed in specialized ways, such as the chewing organ called "
Aristotle's lantern Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) of ...
" in sea urchins, the supportive stalks of crinoids, and the structural "lime ring" of sea cucumbers. Although individual ossicles are robust and fossilize readily, complete skeletons of starfish, brittle stars and crinoids are rare in the fossil record. On the other hand, sea urchins are often well preserved in chalk beds or limestone. During fossilization, the cavities in the stereom are filled in with calcite that is continuous with the surrounding rock. On fracturing such rock,
paleontologists Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...
can observe distinctive cleavage patterns and sometimes even the intricate internal and external structure of the test. The epidermis contains pigment cells that provide the often vivid colours of echinoderms, which include deep red, stripes of black and white, and intense purple. These cells may be light-sensitive, causing many echinoderms to change appearance completely as night falls. The reaction can happen quickly: the sea urchin ''
Centrostephanus longispinus ''Centrostephanus longispinus'', the hatpin urchin, is a species of sea urchin in the family Diadematidae. There are two subspecies, ''Centrostephanus l. longispinus'', found in the eastern Atlantic and Mediterranean Sea and ''Centrostephanus l. ...
'' changes colour in just fifty minutes when exposed to light. One characteristic of most echinoderms is a special kind of tissue known as catch connective tissue. This collagen-based material can change its mechanical properties under nervous control rather than by muscular means. This tissue enables a starfish to go from moving flexibly around the seabed to becoming rigid while prying open a bivalve mollusc or preventing itself from being extracted from a crevice. Similarly, sea urchins can lock their normally mobile spines upright as a defensive mechanism when attacked.


The water vascular system

Echinoderms possess a unique water vascular system, a network of fluid-filled canals derived from the
coelom The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it r ...
(body cavity) that function in gas exchange, feeding, sensory reception and locomotion. This system varies between different classes of echinoderm but typically opens to the exterior through a sieve-like
madreporite The madreporite is a light colored calcareous opening used to filter water into the water vascular system of echinoderms. It acts like a pressure-equalizing valve. It is visible as a small red or yellow button-like structure, looking like a sm ...
on the aboral (upper) surface of the animal. The madreporite is linked to a slender duct, the stone canal, which extends to a ring canal that encircles the mouth or
oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
. The ring canal branches into a set of radial canals, which in asteroids extend along the arms, and in echinoids adjoin the test in the ambulacral areas. Short lateral canals branch off the radial canals, each one ending in an ampulla. Part of the ampulla can protrude through a pore (or a pair of pores in sea urchins) to the exterior, forming a podium or tube foot. The water vascular system assists with the distribution of nutrients throughout the animal's body; it is most visible in the tube feet which can be extended or contracted by the redistribution of fluid between the foot and the internal ampulla. The organisation of the water vascular system is somewhat different in ophiuroids, where the madreporite may be on the oral surface and the podia lack suckers. In holothuroids, the system is reduced, often with few tube feet other than the specialised feeding tentacles, and the madreporite opens on to the coelom. Some holothuroids like the Apodida lack tube feet and canals along the body; others have longitudinal canals. The arrangements in crinoids is similar to that in asteroids, but the tube feet lack suckers and are used in a back-and-forth wafting motion to pass food particles captured by the arms towards the central mouth. In the asteroids, the same motion is employed to move the animal across the ground.


Other organs

Echinoderms possess a simple digestive system which varies according to the animal's diet. Starfish are mostly carnivorous and have a mouth, oesophagus, two-part stomach, intestine and rectum, with the anus located in the centre of the aboral body surface. With a few exceptions, the members of the order Paxillosida do not possess an anus. In many species of starfish, the large cardiac stomach can be everted to digest food outside the body. Some other species are able to ingest whole food items such as molluscs. Brittle stars, which have varying diets, have a blind gut with no intestine or anus; they expel food waste through their mouth. Sea urchins are herbivores and use their specialised mouthparts to graze, tear and chew their food, mainly
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
. They have an oesophagus, a large stomach and a rectum with the anus at the apex of the test. Sea cucumbers are mostly detritivores, sorting through the sediment with modified tube feet around their mouth, the buccal tentacles. Sand and mud accompanies their food through their simple gut, which has a long coiled intestine and a large
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, an ...
. Crinoids are
suspension feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s, passively catching plankton which drift into their outstretched arms. Boluses of mucus-trapped food are passed to the mouth, which is linked to the anus by a loop consisting of a short oesophagus and longer intestine. The coelomic cavities of echinoderms are complex. Aside from the water vascular system, echinoderms have a haemal coelom, a peri
visceral In biology, an organ is a collection of tissues joined in a structural unit to serve a common function. In the hierarchy of life, an organ lies between tissue and an organ system. Tissues are formed from same type cells to act together in a f ...
coelom, a gonadal coelom and often also a perihaemal coelom. During development, echinoderm coelom is divided into the metacoel, mesocoel and protocoel (also called somatocoel, hydrocoel and axocoel, respectively). The water vascular system, haemal system and perihaemal system form the tubular coelomic system. Echinoderms are unusual in having both a coelomic circulatory system (the water vascular system) and a haemal circulatory system, as most groups of animals have just one of the two. Haemal and perihaemal systems are derived from the original coelom, forming an open and reduced circulatory system. This usually consists of a central ring and five radial vessels. There is no true
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide to ...
, and the blood often lacks any respiratory pigment. Gaseous exchange occurs via dermal branchiae or papulae in starfish, genital bursae in brittle stars, peristominal gills in sea urchins and cloacal trees in sea cucumbers. Exchange of gases also takes place through the tube feet. Echinoderms lack specialized excretory (waste disposal) organs and so nitrogenous waste, chiefly in the form of
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...
, diffuses out through the respiratory surfaces. The coelomic fluid contains the
coelomocyte A coelomocyte () is a phagocytic leukocyte that appears in the bodies of animals that have a coelom. In most, it attacks and digests invading organisms such as bacteria and viruses through encapsulation and phagocytosis, though in some animals (e.g. ...
s, or immune cells. There are several types of immune cells, which vary among classes and species. All classes possess a type of phagocytic amebocyte, which engulf invading particles and infected cells, aggregate or clot, and may be involved in cytotoxicity. These cells are usually large and granular, and are believed to be a main line of defense against potential pathogens. Depending on the class, echinoderms may have
spherule A sphere () is a geometrical object that is a three-dimensional analogue to a two-dimensional circle. A sphere is the set of points that are all at the same distance from a given point in three-dimensional space.. That given point is the ...
cells (for cytotoxicity, inflammation, and anti-bacterial activity), vibratile cells (for coelomic fluid movement and clotting), and crystal cells (which may serve for osmoregulation in sea cucumbers). The coelomocytes secrete
antimicrobial peptides Antimicrobial peptides (AMPs), also called host defence peptides (HDPs) are part of the innate immune response found among all classes of life. Fundamental differences exist between prokaryotic and eukaryotic cells that may represent targets for an ...
against bacteria, and have a set of
lectin Lectins are carbohydrate-binding proteins that are highly specific for sugar groups that are part of other molecules, so cause agglutination of particular cells or precipitation of glycoconjugates and polysaccharides. Lectins have a role in r ...
s and
complement proteins The complement system, also known as complement cascade, is a part of the immune system that enhances (complements) the ability of antibodies and phagocytic cells to clear microbes and damaged cells from an organism, promote inflammation, and at ...
as part of an innate immune system that is still being characterised. Echinoderms have a simple radial nervous system that consists of a modified
nerve net A nerve net consists of interconnected neurons lacking a brain or any form of cephalization. While organisms with bilateral body symmetry are normally associated with a condensation of neurons or, in more advanced forms, a central nervous syst ...
of interconnecting neurons with no central brain, although some do possess ganglia. Nerves radiate from central rings around the mouth into each arm or along the body wall; the branches of these nerves coordinate the movements of the organism and the synchronisation of the tube feet. Starfish have sensory cells in the epithelium and have simple eyespots and touch-sensitive tentacle-like tube feet at the tips of their arms. Sea urchins have no particular sense organs but do have
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in ''Xenoturbella''. The statocyst cons ...
s that assist in gravitational orientation, and they have sensory cells in their epidermis, particularly in the tube feet, spines and
pedicellariae A pedicellaria (plural: pedicellariae) is a small wrench- or claw-shaped appendage with movable jaws, called valves, commonly found on echinoderms (phylum Echinodermata), particularly in sea stars (class Asteroidea) and sea urchins (class Echinoi ...
. Brittle stars, crinoids and sea cucumbers in general do not have sensory organs, but some burrowing sea cucumbers of the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagi ...
have a single statocyst adjoining each radial nerve, and some have an eyespot at the base of each tentacle. The gonads at least periodically occupy much of the body cavities of sea urchins and sea cucumbers, while the less voluminous crinoids, brittle stars and starfish have two gonads in each arm. While the ancestors of modern echinoderms are believed to have had one genital aperture, many organisms have multiple gonopores through which eggs or sperm may be released.


Regeneration

Many echinoderms have great powers of regeneration. Many species routinely autotomize and regenerate arms and viscera. Sea cucumbers often discharge parts of their internal organs if they perceive themselves to be threatened, regenerating them over the course of several months. Sea urchins constantly replace spines lost through damage, while sea stars and sea lilies readily lose and regenerate their arms. In most cases, a single severed arm cannot grow into a new starfish in the absence of at least part of the disc.See last paragraph in review above Analysis However, in a few species a single arm can survive and develop into a complete individual, and arms are sometimes intentionally detached for the purpose of
asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the f ...
. During periods when they have lost their digestive tracts, sea cucumbers live off stored nutrients and absorb dissolved organic matter directly from the water. The regeneration of lost parts involves both epimorphosis and morphallaxis. In epimorphosis stem cells—either from a reserve pool or those produced by
dedifferentiation Dedifferentiation (pronounced dē-ˌdi-fə-ˌren-chē-ˈā-shən) is a transient process by which cells become less specialized and return to an earlier cell state within the same lineage. This suggests an increase in a cell potency, meaning that ...
—form a
blastema A blastema (Greek ''βλάστημα'', "offspring") is a mass of cells capable of growth and regeneration into organs or body parts. The changing definition of the word "blastema" has been reviewed by Holland (2021). A broad survey of how blast ...
and generate new tissues. Morphallactic regeneration involves the movement and remodelling of existing tissues to replace lost parts. Direct
transdifferentiation Transdifferentiation, also known as lineage reprogramming, is the process in which one mature somatic cell is transformed into another mature somatic cell without undergoing an intermediate pluripotent state or progenitor cell type. It is a type ...
of one type of tissue to another during tissue replacement is also observed.


Reproduction


Sexual reproduction

Echinoderms become sexually mature after approximately two to three years, depending on the species and the environmental conditions. Almost all species have separate male and female sexes, though some are
hermaphroditic In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have sepa ...
. The eggs and sperm cells are typically released into open water, where fertilisation takes place. The release of sperm and eggs is synchronised in some species, usually with regard to the lunar cycle. In other species, individuals may aggregate during the reproductive season, increasing the likelihood of successful fertilisation. Internal fertilisation has been observed in three species of sea star, three brittle stars and a deep water sea cucumber. Even at abyssal depths, where no light penetrates, echinoderms often synchronise their reproductive activity. Some echinoderms brood their eggs. This is especially common in cold water species where planktonic larvae might not be able to find sufficient food. These retained eggs are usually few in number and are supplied with large yolks to nourish the developing embryos. In starfish, the female may carry the eggs in special pouches, under her arms, under her arched body, or even in her cardiac stomach. Many brittle stars are hermaphrodites; they often brood their eggs, usually in special chambers on their oral surfaces, but sometimes in the ovary or coelom. In these starfish and brittle stars, development is usually direct to the adult form, without passing through a bilateral larval stage. A few sea urchins and one species of sand dollar carry their eggs in cavities, or near their anus, holding them in place with their spines. Some sea cucumbers use their buccal tentacles to transfer their eggs to their underside or back, where they are retained. In a very small number of species, the eggs are retained in the coelom where they develop viviparously, later emerging through ruptures in the body wall. In some crinoids, the embryos develop in special breeding bags, where the eggs are held until sperm released by a male happens to find them.


Asexual reproduction

One species of
seastar Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
, '' Ophidiaster granifer'', reproduces asexually by parthenogenesis. In certain other
asterozoa The Asterozoa are a subphylum in the phylum Echinodermata. Characteristics include a star-shaped body and radially divergent axes of symmetry. The subphylum includes the class Asteroidea (the starfish), the class Ophiuroidea (the brittle stars ...
ns, adults reproduce asexually until they mature, then reproduce sexually. In most of these species, asexual reproduction is by transverse fission with the disc splitting in two. Both the lost disc area and the missing arms regrow, so an individual may have arms of varying lengths. During the period of regrowth, they have a few tiny arms and one large arm, and are thus often known as "comets". Adult sea cucumbers reproduce asexually by transverse fission. '' Holothuria parvula'' uses this method frequently, splitting into two a little in front of the midpoint. The two halves each regenerate their missing organs over a period of several months, but the missing genital organs are often very slow to develop. The larvae of some echinoderms are capable of asexual reproduction. This has long been known to occur among starfish and brittle stars, but has more recently been observed in a sea cucumber, a sand dollar and a sea urchin. This may be by autotomising parts that develop into secondary larvae, by
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is know ...
, or by splitting transversely. Autotomised parts or buds may develop directly into fully formed larvae, or may pass through a gastrula or even a
blastula Blastulation is the stage in early animal embryonic development that produces the blastula. In mammalian development the blastula develops into the blastocyst with a differentiated inner cell mass and an outer trophectoderm. The blastula (fro ...
stage. New larvae can develop from the preoral hood (a mound like structure above the mouth), the side body wall, the postero-lateral arms, or their rear ends. Cloning is costly to the larva both in resources and in development time. Larvae undergo this process when food is plentiful or temperature conditions are optimal. Cloning may occur to make use of the tissues that are normally lost during metamorphosis. The larvae of some sand dollars clone themselves when they detect dissolved fish mucus, indicating the presence of predators. Asexual reproduction produces many smaller larvae that escape better from planktivorous fish, implying that the mechanism may be an anti-predator adaptation.


Larval development

Development begins with a bilaterally symmetrical embryo, with a coeloblastula developing first. Gastrulation marks the opening of the "second mouth" that places echinoderms within the deuterostomes, and the mesoderm, which will host the skeleton, migrates inwards. The secondary body cavity, the coelom, forms by the partitioning of three body cavities. The larvae are often planktonic, but in some species the eggs are retained inside the female, while in some the female broods the larvae. The larvae pass through several stages, which have specific names derived from the taxonomic names of the adults or from their appearance. For example, a sea urchin has an 'echinopluteus' larva while a brittle star has an 'ophiopluteus' larva. A starfish has a '
bipinnaria A bipinnaria is the first stage in the larval development of most starfish, and is usually followed by a brachiolaria stage. Movement and feeding is accomplished by the bands of cilia. Starfish that brood their young generally lack a bipinnaria sta ...
' larva, which develops into a multi-armed '
brachiolaria A brachiolaria is the second stage of larval development in many starfishes. It follows the bipinnaria. Brachiolaria have bilateral symmetry, unlike the adult starfish, which have a pentaradial symmetry. Starfish of the order Paxillosida ('' Astr ...
' larva. A sea cucumber's larva is an 'auricularia' while a crinoid's is a 'vitellaria'. All these larvae are
bilaterally symmetrical Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pl ...
and have bands of cilia with which they swim; some, usually known as 'pluteus' larvae, have arms. When fully developed they settle on the seabed to undergo metamorphosis, and the larval arms and gut degenerate. The left hand side of the larva develops into the oral surface of the juvenile, while the right side becomes the aboral surface. At this stage the pentaradial symmetry develops. A plankton-eating larva, living and feeding in the water column, is considered to be the ancestral larval type for echinoderms, but in extant echinoderms, some 68% of species develop using a yolk-feeding larva. The provision of a yolk-sac means that smaller numbers of eggs are produced, the larvae have a shorter development period and a smaller dispersal potential, but a greater chance of survival.


Distribution and habitat

Echinoderms are globally distributed in almost all depths, latitudes and environments in the ocean. Adults are mainly
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
, living on the seabed, whereas larvae are often pelagic, living as plankton in the open ocean. Some holothuroid adults such as ''
Pelagothuria ''Pelagothuria'' is a genus of sea cucumbers in the family Pelagothuriidae. It is monotypic, being represented by the single species ''Pelagothuria natatrix''. Characteristics This sea cucumber is somewhat unusual in appearance in comparison ...
'' are however pelagic. Some crinoids are pseudo-planktonic, attaching themselves to floating logs and debris, although this behaviour was exercised most extensively in the Paleozoic, before competition from organisms such as barnacles restricted the extent of the behaviour.


Mode of life


Locomotion

Echinoderms primarily use their tube feet to move about, though some sea urchins also use their spines. The tube feet typically have a tip shaped like a suction pad in which a vacuum can be created by contraction of muscles. This combines with some stickiness from the secretion of mucus to provide adhesion. The tube feet contract and relax in waves which move along the adherent surface, and the animal moves slowly along. Brittle stars are the most agile of the echinoderms. Any one of the arms can form the axis of symmetry, pointing either forwards or back. The animal then moves in a co-ordinated way, propelled by the other four arms. During locomotion, the propelling arms can made either snake-like or rowing movements. Starfish move using their tube feet, keeping their arms almost still, including in genera like '' Pycnopodia'' where the arms are flexible. The oral surface is covered with thousands of tube feet which move out of time with each other, but not in a metachronal rhythm; in some way, however, the tube feet are coordinated, as the animal glides steadily along. Some burrowing starfish have points rather than suckers on their tube feet and they are able to "glide" across the seabed at a faster rate. Sea urchins use their tube feet to move around in a similar way to starfish. Some also use their articulated spines to push or lever themselves along or lift their oral surfaces off the substrate. If a sea urchin is overturned, it can extend its tube feet in one ambulacral area far enough to bring them within reach of the substrate and then successively attach feet from the adjoining area until it is righted. Some species bore into rock, usually by grinding away at the surface with their mouthparts. Sea cucumbers are generally sluggish animals. Many can move on the surface of the sea bed or burrow through sand or mud using peristaltic movements; some have short tube feet on their under surface with which they can creep along in the manner of a starfish. Some species drag themselves along using their buccal tentacles, while others manage to swim with peristaltic movements or rhythmic flexing. Many live in cracks, hollows and burrows and hardly move at all. Some deep water species are pelagic and can float in the water with webbed papillae forming sails or fins. The majority of crinoids are motile, but sea lilies are sessile and attached to hard substrates by stalks. Movement in most sea lilies is limited to bending their stems can bend and rolling and unrolling their arms; a few species can relocate themselves on the seabed by crawling. The sea feathers are unattached and usually live in crevices, under corals or inside sponges with their arms the only visible part. Some sea feathers emerge at night and perch themselves on nearby eminences to better exploit food-bearing currents. Many species can "walk" across the seabed, raising their body with the help of their arms, or swim using their arms. Most species of sea feather, however, are largely sedentary, seldom moving far from their chosen place of concealment.


Feeding

The modes of feeding vary greatly between the different echinoderm taxa. Crinoids and some brittle stars tend to be passive filter-feeders, enmeshing suspended particles from passing water. Most sea urchins are grazers; sea cucumbers are deposit feeders; and the majority of starfish are active hunters. Crinoids catch food particles using the tube feet on their outspread pinnules, move them into the ambulacral grooves, wrap them in mucus, and convey them to the mouth using the cilia lining the grooves. The exact dietary requirements of crinoids have been little researched, but in the laboratory they can be fed with diatoms.
Basket star The Euryalida are an order of brittle stars, which includes large species with either branching arms (called "basket stars") or long and curling arms (called "snake stars"). Characteristics Many of the species in this order have characteristi ...
s are suspension feeders, raising their branched arms to collect
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, while other brittle stars use several methods of feeding. Some are suspension feeders, securing food particles with mucus strands, spines or tube feet on their raised arms. Others are scavengers and detritus feeders. Others again are voracious
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other so ...
s and able to lasso their waterborne prey with a sudden encirclement by their flexible arms. The limbs then bend under the disc to transfer the food to the jaws and mouth. Many sea urchins feed on algae, often scraping off the thin layer of algae covering the surfaces of rocks with their specialised mouthparts known as Aristotle's lantern. Other species devour smaller organisms, which they may catch with their tube feet. They may also feed on dead fish and other animal matter. Sand dollars may perform suspension feeding and feed on phytoplankton, detritus, algal pieces and the bacterial layer surrounding grains of sand. Sea cucumbers are often mobile deposit or suspension feeders, using their buccal podia to actively capture food and then stuffing the particles individually into their buccal cavities. Others ingest large quantities of sediment, absorb the organic matter and pass the indigestible mineral particles through their guts. In this way they disturb and process large volumes of substrate, often leaving characteristic ridges of sediment on the sea bed. Some sea cucumbers live infaunally in burrows, anterior-end down and anus on the surface, swallowing sediment and passing it through their gut. Other burrowers live anterior-end up and wait for detritus to fall into the entrances of the burrows or rake in debris from the surface nearby with their buccal podia. Nearly all starfish are detritus feeders or carnivores, though a few are suspension feeders. Small fish landing on the upper surface may be captured by pedicilaria and dead animal matter may be scavenged but the main prey items are living invertebrates, mostly bivalve molluscs. To feed on one of these, the starfish moves over it, attaches its tube feet and exerts pressure on the valves by arching its back. When a small gap between the valves is formed, the starfish inserts part of its stomach into the prey, excretes digestive enzymes and slowly liquefies the soft body parts. As the adductor muscle of the bivalve relaxes, more stomach is inserted and when digestion is complete, the stomach is returned to its usual position in the starfish with its now liquefied bivalve meal inside it. Other starfish evert the stomach to feed on sponges, sea anemones, corals, detritus and algal films.


Antipredator defence

Despite their low nutrition value and the abundance of indigestible calcite, echinoderms are preyed upon by many organisms, including
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
, sharks,
eider duck Eiders () are large seaducks in the genus ''Somateria''. The three extant species all breed in the cooler latitudes of the Northern Hemisphere. The down feathers of eider ducks, and some other ducks and geese, are used to fill pillows and quil ...
s, gulls,
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...
s, gastropod molluscs, other echinoderms, sea otters,
Arctic fox The Arctic fox (''Vulpes lagopus''), also known as the white fox, polar fox, or snow fox, is a small fox native to the Arctic regions of the Northern Hemisphere and common throughout the Arctic tundra biome. It is well adapted to living in co ...
es and humans. Larger starfish prey on smaller ones; the great quantity of eggs and larva that they produce form part of the
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, consumed by many marine creatures. Crinoids, on the other hand, are relatively free from predation. Antipredator defences include the presence of spines, toxins (inherent or delivered through the tube feet), and the discharge of sticky entangling threads by sea cucumbers. Although most echinoderm spines are blunt, those of the
crown-of-thorns starfish The crown-of-thorns starfish (frequently abbreviated to COTS), ''Acanthaster planci'', is a large starfish that preys upon hard, or stony, coral polyps (Scleractinia). The crown-of-thorns starfish receives its name from venomous thorn-like spine ...
are long and sharp and can cause a painful puncture wound as the epithelium covering them contains a toxin. Because of their catch connective tissue, which can change rapidly from a flaccid to a rigid state, echinoderms are very difficult to dislodge from crevices. Some sea cucumbers have a cluster of
cuvierian tubules Cuvierian tubules are clusters of fine tubes located at the base of the respiratory tree in some sea cucumbers in the genera '' Bohadschia'', ''Holothuria'' and '' Pearsonothuria'', all of which are included in the family Holothuriidae. The tubule ...
which can be ejected as long sticky threads from their anus to entangle and permanently disable an attacker. Sea cucumbers occasionally defend themselves by rupturing their body wall and discharging the gut and internal organs. Starfish and brittle stars may undergo
autotomy Autotomy (from the Greek ''auto-'', "self-" and ''tome'', "severing", αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usually as a self-defense mechanism to elude ...
when attacked, detaching an arm; this may distract the predator for long enough for the animal to escape. Some starfish species can swim away from danger.


Ecology

Echinoderms are numerous invertebrates whose adults play an important role in benthic ecosystems, while the larvae are a major component of the plankton. Among the ecological roles of adults are the grazing of sea urchins, the sediment processing of heart urchins, and the suspension and deposit feeding of crinoids and sea cucumbers. Some sea urchins can bore into solid rock, destabilising rock faces and releasing nutrients into the ocean. Coral reefs are also bored into in this way, but the rate of accretion of carbonate material is often greater than the erosion produced by the sea urchin. Echinoderms sequester about 0.1 gigatonnes of carbon dioxide per year as calcium carbonate, making them important contributors in the global
carbon cycle The carbon cycle is the biogeochemical cycle by which carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of the Earth. Carbon is the main component of biological compounds as well as a major componen ...
. Echinoderms sometimes have large population swings which can transform ecosystems. In 1983, for example, the mass mortality of the tropical sea urchin ''
Diadema antillarum ''Diadema antillarum'', also known as the lime urchin, black sea urchin, or the long-spined sea urchin, is a species of sea urchin in the family Diadematidae. This sea urchin is characterized by its exceptionally long black spines. It is the ...
'' in the Caribbean caused a change from a coral-dominated reef system to an alga-dominated one. Sea urchins are among the main herbivores on reefs and there is usually a fine balance between the urchins and the kelp and other algae on which they graze. A diminution of the numbers of predators (otters, lobsters and fish) can result in an increase in urchin numbers, causing overgrazing of kelp forests, resulting in an alga-denuded "
urchin barren An urchin barren is an area of the shallow part of the ocean where the population growth of sea urchins has gone unchecked, causing destructive grazing of kelp forests. Process Sea urchins eat kelp holdfasts. This can be caused by a lack of sea ...
". On the Great Barrier Reef, an unexplained increase in the numbers of
crown-of-thorns starfish The crown-of-thorns starfish (frequently abbreviated to COTS), ''Acanthaster planci'', is a large starfish that preys upon hard, or stony, coral polyps (Scleractinia). The crown-of-thorns starfish receives its name from venomous thorn-like spine ...
(''Acanthaster planci''), which graze on living coral tissue, has greatly increased coral mortality and reduced coral reef
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
.


Use by humans


As food and medicine

In 2019, 129,000 tonnes of echinoderms were harvested. The majority of these were sea cucumbers (158,000 tonnes) and sea urchins (73,000 tonnes). These are used mainly for food, but also in traditional Chinese medicine. Sea cucumbers are considered a delicacy in some countries of southeast Asia; as such, they are in imminent danger of being over-harvested. Popular species include the pineapple roller ''
Thelenota ananas ''Thelenota ananas'', also known as pineapple sea cucumber, oloturia ananas, tripang, prickly skin cucumber, pointed teat sea cucumber, armoured sea cucumber, giant sea cucumber, sand fish or prickly redfish, is a species of sea cucumber found in ...
'' (''susuhan'') and the red sea cucumber ''
Holothuria edulis ''Holothuria edulis'', commonly known as the edible sea cucumber or the pink and black sea cucumber, is a species of echinoderm in the family Holothuriidae. It was placed in the subgenus ''Halodeima'' by Pearson in 1914, making its full scientifi ...
''. These and other species are colloquially known as ''bêche de mer'' or ''trepang'' in China and Indonesia. The sea cucumbers are boiled for twenty minutes and then dried both naturally and later over a fire which gives them a smoky tang. In China they are used as a basis for gelatinous soups and stews. Both male and female gonads of sea urchins are consumed, particularly in Japan and France. The taste is described as soft and melting, like a mixture of seafood and fruit.Davidson, Alan (2014) ''
Oxford Companion to Food ''The Oxford Companion to Food'' is an encyclopedia about food. It was edited by Alan Davidson and published by Oxford University Press in 1999. It was also issued in softcover under the name ''The Penguin Companion to Food''. The second and th ...
''. Oxford University Press, 3rd edition. p. 730.
Sea urchin breeding trials have been undertaken to try to compensate for overexploitation.


In research

Because of their robust larval growth, sea urchins are widely used in research, particularly as model organisms in
developmental biology Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of stem c ...
and ecotoxicology. ''
Strongylocentrotus purpuratus ''Strongylocentrotus purpuratus'', the purple sea urchin, lives along the eastern edge of the Pacific Ocean extending from Ensenada, Mexico, to British Columbia, Canada. This sea urchin species is deep purple in color, and lives in lower int ...
'' and ''
Arbacia punctulata The Atlantic purple sea urchin (''Arbacia punctulata'') is a species of sea urchins from the family Arbaciidae, native to the Atlantic Ocean. Description The Atlantic purple sea urchin is a spherical, dark purple-spined sea urchin, with a ne ...
'' are used for this purpose in embryological studies. The large size and the transparency of the eggs enables the observation of
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, wh ...
cells in the process of fertilising
ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
. The arm regeneration potential of brittle stars is being studied in connection with understanding and treating
neurodegenerative A neurodegenerative disease is caused by the progressive loss of structure or function of neurons, in the process known as neurodegeneration. Such neuronal damage may ultimately involve cell death. Neurodegenerative diseases include amyotrophic ...
diseases in humans.


Other uses

The calcareous tests or shells of echinoderms are used as a source of lime by farmers in areas where limestone is unavailable and some are used in the manufacture of fish meal. Four thousand tons of the animals are used annually for these purposes. This trade is often carried out in conjunction with
shellfish Shellfish is a colloquial and fisheries term for exoskeleton-bearing aquatic invertebrates used as food, including various species of molluscs, crustaceans, and echinoderms. Although most kinds of shellfish are harvested from saltwater enviro ...
farmers, for whom the starfish pose a major threat by eating their cultured stock. Other uses for the starfish they recover include the manufacture of animal feed, composting and the preparation of dried specimens for the arts and craft trade.


See also

*


References


Cited texts

* * *


External links


The Echinoid Directory
from the Natural History Museum
Echinodermata
from the
Tree of Life Web Project The Tree of Life Web Project is an Internet project providing information about the diversity and phylogeny of life on Earth. This collaborative peer reviewed project began in 1995, and is written by biologists from around the world. The site h ...

Echinoderms of the North Sea


{{Authority control * Extant Cambrian first appearances Marine animals