Sea urchins or urchins () are in the class Echinoidea. About 950 species live on the seabed, inhabiting all oceans and depth zones from the intertidal zone to of . They typically have a globular body covered by a spiny protective tests (hard shells), typically from across. Sea urchins move slowly, crawling with their tube feet, and sometimes pushing themselves with their spines. They feed primarily on algae but also eat slow-moving or sessile animals such as and . Their predators include , , starfish, , and triggerfish.
Like all echinoderms, adult sea urchins have pentagonal symmetry with their pluteus larvae featuring bilateral (mirror) symmetry; The latter indicates that they belong to the Bilateria, along with , , and . Sea urchins are found in every ocean and in every climate, from the tropics to the polar regions, and inhabit marine benthic (sea bed) habitats, from rocky shores to hadal zone depths. The fossil record of the echinoids dates from the Ordovician period, some 450 million years ago. The closest echinoderm relatives of the sea urchin are the (Holothuroidea), which like them are , a clade that includes the . ( are a separate order in the sea urchin class Echinoidea.)
The animals have been studied since the 19th century as in developmental biology, as their embryos were easy to observe. That has continued with studies of their because of their unusual fivefold symmetry and relationship to chordates. Species such as the slate pencil urchin are popular in aquaria, where they are useful for controlling algae. Fossil urchins have been used as protective .
Specifically, the term "sea urchin" refers to the "regular echinoids", which are symmetrical and globular, and includes several different taxonomic groups, with two subclasses: Euechinoidea ("modern" sea urchins, including irregular ones) and Cidaroidea, or "slate-pencil urchins", which have very thick, blunt spines, with algae and sponges growing on them. The "irregular" sea urchins are an infra-class inside the Euechinoidea, called Irregularia, and include Atelostomata and Neognathostomata. Irregular echinoids include flattened , Clypeasteroida, and Echinocardium.
Together with sea cucumbers (Holothuroidea), they make up the subphylum Echinozoa, which is characterized by a globoid shape without arms or projecting rays. Sea cucumbers and the irregular echinoids have secondarily evolved diverse shapes. Although many sea cucumbers have branched surrounding their oral openings, these have originated from modified tube feet and are not homologous to the arms of the crinoids, sea stars, and brittle stars.
Like other echinoderms, sea urchin early larvae have bilateral symmetry,Stachan and Read, Human Molecular Genetics, "What Makes Us Human", p. 381. but they develop five-fold symmetry as they mature. This is most apparent in the "regular" sea urchins, which have roughly spherical bodies with five equally sized parts radiating out from their central axes. The mouth is at the base of the animal and the anus at the top; the lower surface is described as "oral" and the upper surface as "aboral".
Several sea urchins, however, including the sand dollars, are oval in shape, with distinct front and rear ends, giving them a degree of bilateral symmetry. In these urchins, the upper surface of the body is slightly domed, but the underside is flat, while the sides are devoid of tube feet. This "irregular" body form has evolved to allow the animals to burrow through sand or other soft materials.
The test is rigid, and divides into five ambulacral grooves separated by five wider interambulacral areas. Each of these ten longitudinal columns consists of two sets of plates (thus comprising 20 columns in total). The ambulacral plates have pairs of tiny holes through which the tube feet extend.
All of the plates are covered in rounded tubercles to which the spines are attached. The spines are used for defence and for locomotion and come in a variety of forms. The inner surface of the test is lined by peritoneum. Sea urchins convert aqueous carbon dioxide using a catalytic process involving nickel into the calcium carbonate portion of the test.
Most species have two series of spines, primary (long) and secondary (short), distributed over the surface of the body, with the shortest at the poles and the longest at the equator. The spines are usually hollow and cylindrical. Contraction of the muscular sheath that covers the test causes the spines to lean in one direction or another, while an inner sheath of collagen fibres can reversibly change from soft to rigid which can lock the spine in one position. Located among the spines are several types of pedicellaria, moveable stalked structures with jaws.
Sea urchins move by walking, using their many flexible tube feet in a way similar to that of starfish; regular sea urchins do not have any favourite walking direction.Kazuya Yoshimura, Tomoaki Iketani et Tatsuo Motokawa, "Do regular sea urchins show preference in which part of the body they orient forward in their walk ?", Marine Biology, vol. 159, no 5, 2012, p. 959–965. The tube feet protrude through pairs of pores in the test, and are operated by a water vascular system; this works through hydraulic pressure, allowing the sea urchin to pump water into and out of the tube feet. During locomotion, the tube feet are assisted by the spines which can be used for pushing the body along or to lift the test off the substrate. Movement is generally related to feeding, with the red sea urchin ( Mesocentrotus franciscanus) managing about a day when there is ample food, and up to a day where there is not. An inverted sea urchin can right itself by progressively attaching and detaching its tube feet and manipulating its spines to roll its body upright. Some species bury themselves in soft sediment using their spines, and Paracentrotus lividus uses its jaws to burrow into soft rocks.
On the upper surface of the test at the aboral pole is a membrane, the periproct, which surrounds the anus. The periproct contains a variable number of hard plates, five of which, the genital plates, contain the gonopores, and one is modified to contain the madreporite, which is used to balance the water vascular system.
The mouth of most sea urchins is made up of five calcium carbonate teeth or plates, with a fleshy, tongue-like structure within. The entire chewing organ is known as Aristotle's lantern from Aristotle's description in his History of Animals (translated by D'Arcy Thompson):
However, this has recently been proven to be a mistranslation. Aristotle's lantern is actually referring to the whole shape of sea urchins, which look like the ancient lamps of Aristotle's time.
are unusual in not having a lantern. Instead, the mouth is surrounded by cilia that pull strings of mucus containing food particles towards a series of grooves around the mouth.
The lantern, where present, surrounds both the mouth cavity and the pharynx. At the top of the lantern, the pharynx opens into the esophagus, which runs back down the outside of the lantern, to join the small intestine and a single caecum. The small intestine runs in a full circle around the inside of the test, before joining the large intestine, which completes another circuit in the opposite direction. From the large intestine, a rectum ascends towards the anus. Despite the names, the small and large intestines of sea urchins are in no way homologous to the similarly named structures in vertebrates.
Digestion occurs in the intestine, with the caecum producing further digestive . An additional tube, called the siphon, runs beside much of the intestine, opening into it at both ends. It may be involved in resorption of water from food.
Sea urchins possess a hemal system with a complex network of vessels in the mesenteries around the gut, but little is known of the functioning of this system. However, the main circulatory fluid fills the general body cavity, or coelom. This coelomic fluid contains phagocyte coelomocytes, which move through the vascular and hemal systems and are involved in internal transport and gas exchange. The coelomocytes are an essential part of blood clotting, but also collect waste products and actively remove them from the body through the gills and tube feet.
Most sea urchins possess five pairs of external gills attached to the peristomial membrane around their mouths. These thin-walled projections of the body cavity are the main organs of respiration in those urchins that possess them. Fluid can be pumped through the gills' interiors by muscles associated with the lantern, but this does not provide a continuous flow, and occurs only when the animal is low in oxygen. Tube feet can also act as respiratory organs, and are the primary sites of gas exchange in heart urchins and sand dollars, both of which lack gills. The inside of each tube foot is divided by a septum which reduces diffusion between the incoming and outgoing streams of fluid.
Sea urchins are sensitive to touch, light, and chemicals. There are numerous sensitive cells in the epithelium, especially in the spines, pedicellaria and tube feet, and around the mouth. Although they do not have eyes or eye spots (except for Diadematidae, which can follow a threat with their spines), the entire body of most regular sea urchins might function as a compound eye.
An unusual feature of sea urchin development is the replacement of the larva's bilateral symmetry by the adult's broadly fivefold symmetry. During cleavage, mesoderm and small micromeres are specified. At the end of gastrulation, cells of these two types form pouches. In the larval stages, the adult rudiment grows from the left coelomic pouch; after metamorphosis, that rudiment grows to become the adult. The animal-vegetal axis is established before the egg is fertilized. The oral-aboral axis is specified early in cleavage, and the left-right axis appears at the late gastrula stage.
Several months are needed for the larva to complete its development, the change into the adult form beginning with the formation of test plates in a juvenile rudiment which develops on the left side of the larva, its axis being perpendicular to that of the larva. Soon, the larva sinks to the bottom and metamorphosis into a juvenile urchin in as little as one hour. In some species, adults reach their maximum size in about five years. The purple urchin becomes sexually mature in two years and may live for twenty.
Adult sea urchins are usually well protected against most predators by their strong and sharp spines, which can be venomous in some species. The small urchin clingfish lives among the spines of urchins such as Diadema; juveniles feed on the pedicellariae and sphaeridia, adult males choose the tube feet and adult females move away to feed on shrimp eggs and molluscs.
Sea urchins are one of the favourite foods of many , , triggerfish, California sheephead, sea otter and Anarhichadidae (which specialise in sea urchins). All these animals carry particular adaptations (teeth, pincers, claws) and a strength that allow them to overcome the excellent protective features of sea urchins. Left unchecked by predators, urchins devastate their environments, creating what biologists call an urchin barren, devoid of macroalgae and associated fauna. Sea urchins graze on the lower stems of kelp, causing the kelp to drift away and die. Loss of the habitat and nutrients provided by leads to profound cascade effects on the marine ecosystem. Sea otters have re-entered British Columbia, dramatically improving coastal ecosystem health.
are a good means of defense against ectoparasites, but not a panacea as some of them actually feed on it.Hiroko Sakashita, " Sexual dimorphism and food habits of the clingfish, Diademichthys lineatus, and its dependence on host sea urchin ", Environmental Biology of Fishes, vol. 34, no 1, 1994, p. 95–101 The hemal system defends against endoparasites.
Population densities vary by habitat, with more dense populations in barren areas as compared to kelp stands. Even in these barren areas, greatest densities are found in shallow water. Populations are generally found in deeper water if wave action is present. Densities decrease in winter when storms cause them to seek protection in cracks and around larger underwater structures. The shingle urchin ( Colobocentrotus atratus), which lives on exposed shorelines, is particularly resistant to wave action. It is one of the few sea urchin that can survive many hours out of water.
Sea urchins can be found in all climates, from warm seas to polar oceans. The larvae of the polar sea urchin Sterechinus neumayeri have been found to use energy in metabolic processes twenty-five times more efficiently than do most other organisms. Antarctic Sea Urchin Shows Amazing Energy-Efficiency in Nature's Deep Freeze 15 March 2001 University of Delaware. Retrieved 22 March 2018 Despite their presence in nearly all the marine ecosystems, most species are found on temperate and tropical coasts, between the surface and some tens of meters deep, close to photosynthetic food sources.
Most fossil echinoids from the Paleozoic era are incomplete, consisting of isolated spines and small clusters of scattered plates from crushed individuals, mostly in Devonian and Carboniferous rocks. The shallow-water from the Ordovician and Silurian periods of Estonia are famous for echinoids. Paleozoic echinoids probably inhabited relatively quiet waters. Because of their thin tests, they would certainly not have survived in the wave-battered coastal waters inhabited by many modern echinoids. Echinoids declined to near extinction at the end of the Paleozoic era, with just six species known from the Permian period. Only two lineages survived this period's massive extinction and into the Triassic: the genus Miocidaris, which gave rise to modern cidaroida (pencil urchins), and the ancestor that gave rise to the euechinoidea. By the upper Triassic, their numbers increased again. Cidaroids have changed very little since the Late Triassic, and are the only Paleozoic echinoid group to have survived.
The euechinoids diversified into new lineages in the Jurassic and Cretaceous periods, and from them emerged the first irregular echinoids (the Atelostomata) during the early Jurassic.
Some echinoids, such as Micraster in the chalk of the Cretaceous period, serve as zone or Index fossil fossils. Because they are abundant and evolved rapidly, they enable geologists to date the surrounding rocks.
In the Paleogene and Neogene periods ( circa 66 to 2.6 Mya), (Clypeasteroida) arose. Their distinctive, flattened tests and tiny spines were adapted to life on or under loose sand in shallow water, and they are abundant as fossils in southern European limestones and sandstones.
The phylogenetic study from 2022 presents a different topology of the Euechinoidea phylogenetic tree. Irregularia are sister group of Echinacea (including Salenioida) forming a common clade Carinacea, basal groups Aspidodiadematoida, Diadematoida, Echinothurioida, Micropygoida, and Pedinoida are comprised in a common basal clade Aulodonta.
The organism's evolutionary placement and unique embryology with five-fold symmetry were the major arguments in the proposal to seek the sequencing of its genome. Importantly, urchins act as the closest living relative to chordates and thus are of interest for the light they can shed on the evolution of . The genome of Strongylocentrotus purpuratus was completed in 2006 and established homology between sea urchin and vertebrate immune system-related genes. Sea urchins code for at least 222 Toll-like receptor genes and over 200 genes related to the Nod-like-receptor family found in vertebrates. This increases its usefulness as a valuable model organism for studying the evolution of innate immunity. The sequencing also revealed that while some genes were thought to be limited to vertebrates, there were also innovations that have previously never been seen outside the chordate classification, such as immune transcription factors PU.1 and SPIB.
Sea urchins are commonly eaten stuffed with rice in the traditional oko-oko dish among the Sama-Bajau people of the Philippines. They were once foraged by coastal Malay communities of Singapore who call them jani. In New Zealand, Evechinus chloroticus, known as kina in Māori, is a delicacy, traditionally eaten raw. Though New Zealand fishermen would like to export them to Japan, their quality is too variable.
In Mediterranean cuisines, Paracentrotus lividus is often eaten raw, or with lemon,for Puglia, Italy: Touring Club Italiano, Guida all'Italia gastronomica, 1984, p. 314; for Alexandria, Egypt: Claudia Roden, A Book of Middle Eastern Food, p. 183 and known as ricci on Italian menus where it is sometimes used in pasta sauces. It can also flavour , scrambled eggs, fish soup,Alan Davidson, Mediterranean Seafood, p. 270 mayonnaise, béchamel sauce for tartlets,Larousse Gastronomique the boullie for a soufflé,Curnonsky, Cuisine et vins de France, nouvelle édition, 1974, p. 248 or Hollandaise sauce to make a fish sauce.Davidson, Alan (2014) Oxford Companion to Food. Oxford University Press, 3rd edition. p. 280 In the region of Marseille, sea urchin are commonly eaten in dedicated food festival called oursinade.Every year, 'oursinades' (sea urchin festivals) are held in and around Marseille to promote this '
On the Pacific Coast of North America, Strongylocentrotus franciscanus was praised by Euell Gibbons; Strongylocentrotus purpuratus is also eaten. Native Americans in California are also known to eat sea urchins. The coast of Southern California is known as a source of high quality , with divers picking sea urchin from kelp beds in depths as deep as 24 m/80 ft. As of 2013, the state was limiting the practice to 300 sea urchin diver licenses. Though the edible Strongylocentrotus droebachiensis is found in the North Atlantic, it is not widely eaten. However, sea urchins (called uutuk in Alutiiq language) are commonly eaten by the Alaska Native population around Kodiak Island. It is commonly exported, mostly to Japan. In the West Indies, slate pencil urchins are eaten.Davidson, Alan (2014) Oxford Companion to Food. Oxford University Press, 3rd edition. pp. 730–731.
In Chilean cuisine, it is served raw with lemon, onions, and olive oil.
Description
Systems
Musculoskeletal
Feeding and digestion
Circulation and respiration
Nervous system and senses
In general, sea urchins are negatively attracted to light, and seek to hide themselves in crevices or under objects. Most species, apart from Cidaris, have in globular organs called spheridia. These are stalked structures and are located within the ambulacral areas; their function is to help in gravitational orientation.
Life history
Reproduction
Development
Life cycle and development
Longevity
Ecology
Trophic level
Predators, parasites, and diseases
Anti-predator defences
Range and habitat
Evolution
Fossil history
Phylogeny
External
Internal
Relation to humans
Injuries
Science
As food
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Aquaria
Folklore
Explanatory notes
External links
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