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Holothuroidea
A sea cucumber
Scientific classification
Domain: Eukarya
Kingdom: Animalia
Phylum: Echinodermata
Subphylum: Echinozoa
Class: Holothuroidea
de Blainville
Orders

Holothuroidea are a class of marine animals (phylum Echinodermata) with an elongated body and leathery skin, which is found on the sea floor worldwide. Many holothurian species and genera, informally known as sea cucumbers, are targeted for human consumption. The harvested product is also referred to as sea cucumber, or as trepang, bêche-de-mer, balate, or sea slug. The body contains a single, branched gonad.

Like all echinoderms, sea cucumbers have an endoskeleton just below the skin, calcified structures that are usually reduced to isolated microscopic ossicles (or sclerietes) joined by connective tissue. These can sometimes be enlarged to flattened plates, forming an armour. In pelagic species such as Pelagothuria natatrix (Order Elasipodida, family Pelagothuriidae), the skeleton and a calcareous ring are absent[1][2].

Contents

Overview

Conspicuous Sea Cucumber, Coconut Island, Hawaii
A sea cucumber feeding while on gravel
Sea cucumber in Mahé, Seychelles ejects sticky filaments from the anus in self-defence.

Sea cucumbers communicate with each other through sending hormone signals through the water which others pick up.

A remarkable feature of these animals is the catch collagen that forms their body wall. This can be loosened and tightened at will, and if the animal wants to squeeze through a small gap, it can essentially liquefy its body and pour into the space. To keep itself safe in these crevices and cracks, the sea cucumber hooks up all its collagen fibres to make its body firm again.[3]

Some species of coral-reef sea cucumbers within the order Aspidochirotida can defend themselves by expelling their sticky cuvierian tubules (enlargements of the respiratory tree that float freely in the coelom) to entangle potential predators. When startled, these cucumbers may expel some of them through a tear in the wall of the cloaca in an autotomic process known as evisceration. Replacement tubules grow back in one-and-a-half to five weeks, depending on the species.[4] The release of these tubules can also be accompanied by the discharge of a toxic chemical known as holothurin, which has similar properties to soap. This chemical can kill any animal in the vicinity and is one more way in which these sedentary animals can defend themselves. [3]

They can be found in great numbers on the deep sea floor, where they often make up the majority of the animal biomass.[5] At depths deeper than 5.5 mi (8.8 km), sea cucumbers comprise 90% of the total mass of the macrofauna[6]. Sea Cucumbers form large herds that move across the bathygraphic features of the ocean, hunting food. The body of some deep water holothurians is made of a tough gelatinous tissue with unique properties that makes the animals able to control their own buoyancy, making it possible for them to either live on the ocean floor or to float over it to move to new locations with a minimum of energy.[7], for instance Enypniastes eximia, Peniagone leander and Paelopatides confundens[8].

In more shallow waters, sea cucumbers can form dense populations. The strawberry sea cucumber (Squamocnus brevidentis) of New Zealand lives on rocky walls around the southern coast of the South Island where populations sometimes reach densities of 1,000 animals per square metre. For this reason, one such area in Fiordland is simply called the strawberry fields.[9]

A variety of fish, most commonly pearl fish, have evolved a commensalistic symbiotic relationship with sea cucumbers in which the pearl fish will live in sea cucumber's cloaca using it for protection from predation, a source of food (the nutrients passing in and out of the anus from the water), and to develop into their adult stage of life.

Emperor shrimp Periclimenes imperator on a Bohadschia argus sea cucumber

Many polychaete worms and crabs have also specialized to use the cloacal respiratory trees for protection by living inside the sea cucumber.[10]

The largest American species is Holothuria floridana, which abounds just below low-water mark on the Florida reefs.

Visitors to Guam often encounter the local variation, called balate, which litters the sea floor all around the island, including in water as shallow as 3 feet (91 cm). These jet black sea cucumbers are normally 10 to 12 inches (25 to 30 cm) long, 1.5 to 2.0 inches (3.8 to 5.1 cm) in diameter and are often curled up, partially covered with sand from the sea floor.

The most common way to separate the subclasses is by looking at their oral tentacles. Subclass Dendrochirotacea has 8-30 oral tentacles, subclass Aspidochirotacea has 10-30 leaflike or shieldlike oral tentacles, while subclass Apodacea may have up to 25 simple or pinnate oral tentacles and is also characterized by reduced or absent tube feet, as in the order Apodida.

Anatomy

Sea cucumbers are typically 10 to 30 centimetres (3.9 to 12 in) in length, although the smallest known species is just 3 millimetres (0.12 in) long, and the largest can reach 1 metre (3.3 ft). The body ranges from almost spherical to worm-like, and lacks the arms found in many other echinoderms, such as starfishes. The anterior end of the animal, containing the mouth, corresponds to the oral pole of other echinoderms (which, in most cases, is the underside), while the posterior end, containing the anus, corresponds to the aboral pole. Thus, compared with other echinoderms, sea cucumbers can be said to be lying on their side.[11]

Like all echinoderms, sea cucumbers possess pentaradial symmetry. However, because of their posture, they have secondarily evolved a degree of bilateral symmetry. For example, because one side of the body is typically pressed against the substratum, and the other is not, there is usually some difference between the two surfaces. Like sea urchins, most sea cucumbers have five strip-like ambulacral areas running along the length of the body from the mouth to the anus. The three on the lower surface have numerous tube feet, often with suckers, that allow the animal to crawl along. The two on the upper surface have under-developed or vestigial tube feet, and, in some species, lack tube feet altogether.[11]

In some species, the ambulacral areas can no longer be distinguished, with tube feet spread over a much wider area of the body. Those of the subclass Apodacea have no tube feet or ambulacral areas at all, and burrow through sediment with muscular contractions of their body.[11]

However, even in those sea cucumbers that lack regular tube feet, those immediately around the mouth are always present. These are highly modified into retractile tentacles, much larger than the regular tube feet. Sea cucumbers have between ten and thirty such tentacles, depending on the species.[11]

Echinoderms typically possess an internal skeleton composed of plates of calcium carbonate. In most sea cucumbers, however, these have become reduced to microscopic ossicles embedded beneath the skin. A few genera, such as Sphaerothuria, retain relatively large plates, giving them a scaly armour.[11]

Diet and digestive system

Holothuroidea are generally scavengers, feeding on debris in the benthic zone of the ocean. Exceptions include pelagic cucumbers and the species Rynkatropa pawsoni, which has a commensal relationship with deep-sea anglerfish.[12] The diet of most cucumbers consists of plankton and decaying organic matter found in the sea. Some sea cucumbers position themselves in currents and catch food that flows by with their open tentacles. They also sift through the bottom sediments using their tentacles.

A pharynx lies behind the mouth and is surrounded by a ring of ten calcareous plates. In most sea cucumbers, this is the only substantial part of the skeleton, and it forms the point of attachment for muscles that can retract the tentacles into the body for safety as for the main muscles of the body wall. Many species possess an oesophagus and stomach, but in some the pharynx opens directly into the intestine. The intestine is typically long and coiled, and loops through the body three times before terminating in a cloacal chamber, or directly as the anus.[11]

Respiratory system

Sea cucumbers extract oxygen from water in a pair of 'respiratory trees' that branch off the cloaca just inside the anus, so that they 'breathe' by drawing water in through the anus and then expelling it.[13][14] The trees consist of a series of narrow tubules branching from a common duct, and lie on either side of the digestive tract. Gas exchange occurs across the thin walls of the tubules, to and from the fluid of the main body cavity.

Together with the intestine, the respiratory trees also act as excretory organs, with nitrogenous waste diffusing across the tubule walls in the form of ammonia and phagocytic coelomocytes depositing particulate waste.[11]

Circulatory systems

Like all echinoderms, sea cucumbers possess both a water vascular system that provides hydraulic pressure to the tentacles and tube feet, allowing them to move, and a haemal system. The latter is more complex than that in other echinoderms, and consists of well-developed vessels as well as open sinuses.[11]

A central haemal ring surrounds the pharynx next to the ring canal of the water vascular system, and sends off additional vessels along the radial canals beneah the ambulacral areas. In the larger species, additional vessels run above and below the intestine and are connected by over a hundred small muscular ampullae, acting as miniature hearts to pump blood around the haemal system. Additional vessels surround the respiratory trees, although they contact them only indirectly, via the coelomic fluid.[11]

Indeed, the blood itself is essentially identical with the coelomic fluid that bathes the organs directly, and also fills the water vascular system. Phagocytic coelomocytes, somewhat similar in function to the white blood cells of vertebrates, are formed within the haemal vessels, and travel throughout the body cavity as well as both circulatory systems. An additional form of coelomocyte, not found in other echinoderms, has a flattened discoid shape, and contains haemoglobin. As a result, in many (though not all) species, both the blood and the coelomic fluid are red in colour.[11]

Ten percent of the blood cell pigment of the sea cucumber is vanadium. Just as the horseshoe crab has blue blood rather than red blood (colored by iron in hemoglobin) because of copper in the hemocyanin pigment, the blood of some sea cucumbers is yellow because of the vanadium in the vanabin pigment[15]. Nonetheless, there is no evidence that vanabins carry oxygen, in contrast to hemoglobin and hemocyanin.

Nervous system

Sea cucumbers have no true brain. A ring of neural tissue surrounds the oral cavity, and sends nerves to the tentacles and the pharynx. The animal is, however, quite capable of functioning and moving about if the nerve ring is surgically removed, demonstrating that it does not have a central role in nervous coordination. In addition, five major nerves run from the nerve ring down length of the body beneath each of the ambulacral areas.[11]

Most sea cucumbers have no distinct sensory organs, although there are various nerve endings scattered through the skin giving the animal a sense of touch and a sensitivity to the presence of light. There are, however, a few exceptions; members of the Apodida order are known to possess statocysts, while some species possess small eye-spots near the bases of their tentacles.[11]

Reproduction and life cycle

Most sea cucumbers reproduce by releasing sperm and ova into the ocean water. Depending on conditions, one organism can produce thousands of gametes. Sea cucumbers are typically dioecious, with separate male and female individuals, but some species are protandric. The reproductive system consists of a single gonad, consisting of a cluster of tubules emptying into a single duct that opens on the upper surface of the animal, close to the tentacles.[11]

At least 30 species, including the red-chested sea cucumber (Pseudocnella insolens), fertilise their eggs internally and then pick up the fertilised zygote with one of their feeding tentacles. The egg is then inserted into a pouch on the adult's body, where it develops and eventually hatches from the pouch as a juvenile sea cucumber[16]. A few species are known to brood their young inside the body cavity, giving birth through a small rupture in the body wall close to the anus.[11]

In all other species, the egg develops into a free-swimming larva, typically after around three days of development. The first stage of larval development is known as an auricularia, and is only around 1 millimetre (0.039 in) in length. This larva swims by means of a long band of cilia wrapped around its body, and somewhat resembles the bipinnaria larva of starfish. As the larva grows it transforms into the doliolaria, with a barrel-shaped body and three to five separate rings of cilia. The tentacles are usually the first adult features to appear, before the regular tube feet.[11]

Holothurians as food and medicine

Dried sea cucumbers in a Japanese pharmacy

To supply the markets of Southern China, Macassan trepangers traded with the Indigenous Australians of Arnhem Land. This Macassan contact with Australia is the first recorded example of trade between the inhabitants of the Australian continent and their Asian neighbours.[17]

There are many of commercially important species of sea cucumber that are harvested and dried for export for use in Chinese cuisine as Hoi sam. Some of the more commonly found species in markets include[18]

Some varieties of sea cucumber (known as gamat in Malaysia or teripang in Indonesia) are said to have excellent healing properties. There are pharmaceutical companies being built based on this gamat product. Extracts are prepared and made into oil, cream, or cosmetics. Some products are intended to be taken internally. The effectiveness of sea cucumber extract in tissue repair has been the subject of serious study.[19] It is believed that the sea cucumber contains all the fatty acids necessary to play an active role in tissue repair.[20]

On December 21, 2007, a study published in PLoS Pathogens found that a lectin from Cucumaria echinata impaired the development of the malaria parasite when produced by transgenic mosquitoes.[21] they are also a chinese tradition to eat

Commercial harvest

In recent years the sea cucumber industry in Alaska has gained strength due to increased export of the skins and muscles to China.[22]

In China, many commercial sea cucumbers are farmed in artificial ponds. These ponds can be as large as 1,000 acres, and satisfy much of the local demand.[22] Wild sea cucumbers are caught by divers and these wild Alaskan sea cucumbers have higher nutritional value and are larger than farmed Chinese sea cucumbers. Larger size and higher nutritional value has allowed the Alaskan fisheries to continue to compete for market shares, despite the increase in local, Chinese sea cucumber farming.[22]

In art and literature

Sea cucumber (a - Tentacles, b - Cloaca, c - Ambulacral feet on the ventral side, d - Papillae on the back)

Sea cucumbers have inspired musical composition: in the first of his Embryons desséchés for piano solo, Erik Satie presents the "(Desiccated embryo) of a Holothurian" and inserts a description of the animal in the score:

The Holothurian crawls across boulders and rocky surfaces.
This sea-animal purrs like a cat; also, it produces disgusting silky threads.
Light appears to have an incommodating effect on it.

Nonetheless it is the sea cucumber's closest relative (the echinoidea or sea urchin) that gets the most attention from scientists, both as an embryo and as a fossil.

Sea cucumbers have also inspired thousands of haiku in Japan, where they are called namako (海鼠), written with characters that can be translated "sea mice". In English translations of these haiku, they are usually called "sea slugs"; according to the Oxford English Dictionary, the term "sea slug" originally referred to holothurians (in the 18th century), though biologists now use the name only for the nudibranch molluscs, marine relatives of land slugs. Almost 1,000 Japanese holothurian haiku translated into English appear in the book Rise, Ye Sea Slugs! by Robin D. Gill (Paraverse Press, 2003, ISBN 0-9742618-0-7).

Refered to as biche de mer, the sea cucumber is mentioned several times in Edgar Allen Poe's only [complete] novel "The Narrative of Arthur Gordon Pym of Nantucket" and a several paragraph description of their biology and preparation as food is given.

Captivity

Sea cucumbers are very common in marine reef aquaria, particularly in reef tanks, where they are prized for their unusual appearance and behavior. Care of sea cucumbers is not complex, but these unusual creatures have unusual requirements. In the hobby, the term sea cucumber refers to only the detrivore sea cucumbers, that is, those that subsist by consuming the detritus that accumulates on the substrate (such as sand or aragonite).

In particular, these creatures have the remarkable ability to live for months, often up to half a year, without feeding. It is very common for these creatures to be introduced into a system that can't support them, and for the owners to have no idea that they are slowly starving to death. When this happens, the sea cucumber will slowly shrink as it digests its own body mass to survive.

In order to be sure a cucumber is feeding one must watch it at work. It will use the feeding tentacles around its mouth to pick up and swallow sand from the bottom of the aquarium. Particles too big will be of no use to the cucumber, so it is important to watch it to make sure it's feeding, and that it's regularly producing castings of excreted substrate.

In addition to the unusual feeding requirements of sea cucumbers, they release highly toxic compounds when injured. In particular, the filter feeding sea cucumbers, known as "Sea Apples" in the aquarium trade, are exceedingly lethal to the other tank inhabitants should they be injured. All powerheads and pumps should be covered as the cucumbers can squeeze into spaces much smaller than their body. Should a sea apple become injured it must be immediately removed from the aquarium, a major water change needs to be performed, and fresh activated carbon will need to be added if there is to be any hope of saving the other inhabitants.

Gallery

See also

Notes

  1. ^ Pelagic sea cucumber: Information from Answers.com
  2. ^ Reich, Mike (30-31 January 2006). Lefebvre, B.; David, B.; Nardin, E. & Poty, E.. ed. "Cambrian holothurians ? – The early fossil record and evolution of Holothuroidea". Journées Georges Ubaghs (Dijon, France: Université de Bourgogne): 36–37. http://www.geobiologie.uni-goettingen.de/people/mreich/pdf/PDFs/POST_Dijon_Seegurken1.pdf.  
  3. ^ a b Piper, Ross (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Greenwood Press. ISBN 0313339228.  
  4. ^ Flammang, Patrick; Ribesse, Jerome & Jangoux, Michel (2002-12-01). "Biomechanics of adhesion in sea cucumber cuvierian tubules (echinodermata, holothuroidea)". Integrative and Comparative Biology 42 (6): 1107–1115. doi:10.1093/icb/42.6.1107. http://icb.oxfordjournals.org/cgi/content/abstract/42/6/1107. Retrieved 2007-10-03.  
  5. ^ Miller, Nat. "Sea Cucumbers". http://jrscience.wcp.muohio.edu/fieldcourses05/PapersMarineEcologyArticles/SeaCucumbers.html. Retrieved 2007-10-03.  
  6. ^ Sci-Tech Encyclopedia: Holothuroidea
  7. ^ Carney, Bob (2007-06-18). "The Kingdom of the Echinoderm". http://www.oceanexplorer.noaa.gov/explorations/07mexico/logs/june18/june18.html. Retrieved 2007-10-03.  
  8. ^ Deep-sea demersal fish zone
  9. ^ Alcock, Niki (2003). "Shedding new light on the humble sea cucumber". Aquatic Biodiversity & Biosecurity Update (New Zealand: National Institute of Water and Atmospheric Research) (3). http://www.niwa.co.nz/news-and-publications/publications/all/abb/2003-03/cucumber. Retrieved 2007-10-03.  
  10. ^ Toonen, Rob, Ph.D. (March 2003). "Aquarium Invertebrates". Advanced Aquarist's Online Magazine 2 (3). http://www.advancedaquarist.com/issues/mar2003/invert.htm. Retrieved 2007-10-03.  
  11. ^ a b c d e f g h i j k l m n o Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 981–997. ISBN 0-03-056747-5.  
  12. ^ Brusca, R.C.; Brusca, G.J. (1990). Invertebrates. Massachusetts: Sinauer Associates. ISBN 0–87893–097–3.  
  13. ^ "Holothurians or sea cucumbers". http://www.fegi.ru/prim/sea/golot.htm. Retrieved 2007-10-03.  
  14. ^ Ingram, Jocie (2006-06-16). "Knowing Nature... Cool as a Sea Cucumber". http://www.comoxvalleynaturalist.bc.ca/knowing_nature/2006/sea_cucumbers.html. Retrieved 2007-10-03.  
  15. ^ Natkin, Michael (2007). "Blood Color". Science Facts. Soak (Source Of All Knowledge). http://www.soak.com/topic/sciencefacts/article/tshow/98556/blood+color. Retrieved 2007-11-16.  
  16. ^ Branch GM, Griffiths CL, Branch ML and Beckley LE (2005) Two Oceans ISBN 0-86486-672-0
  17. ^ Tippett, A. R.. The Voyage to Marege: Macassan Trepangers in Northern Australia.  
  18. ^ Ramofafia C., Byrne M., Battaglene S. C (2003). "Development of three commercial sea cucumbers, Holothuria scabra, H. fuscogilva and Actinopyga mauritiana: larval structure and growth". Marine and freshwater research 54 (5): 657–667. doi:10.1071/MF02145. ISSN 1323-1650.  
  19. ^ B.H. Ridzwan , T.C. Leong and S.Z. Idid (2003). "The Antinociceptive Effects of Water Extracts from Sea Cucumbers Holothuria leucospilota Brandt, Bohadschia marmorata vitiensis Jaeger and Coelomic Fluid from Stichopus hermanii". Pakistan Journal of Biological Sciences 6 (24): 2068–2072. doi:10.3923/pjbs.2003.2068.2072.  
  20. ^ B. D. Fredalina, B. H. Ridzwan, A. A. Zainal Abidin, M. A. Kaswandi, H. Zaiton, I. Zali, P. Kittakoop and A. M. Mat Jais (October 1999). "Fatty acid compositions in local sea cucumber, Stichopus chloronotus, for wound healing". General pharmacology 33 (4): 337–340. doi:10.1016/S0306-3623(98)00253-5. PMID 10523072.  
  21. ^ Yoshida S, Shimada Y, Kondoh D, et al. (2007). "Hemolytic C-type lectin CEL-III from sea cucumber expressed in transgenic mosquitoes impairs malaria parasite development". PLoS Pathog. 3 (12): e192. doi:10.1371/journal.ppat.0030192. PMID 18159942. PMC 2151087. http://www.plospathogens.org/article/info:doi/10.1371/journal.ppat.0030192.  
  22. ^ a b c Ess, Charlie. "Wild product’s versatility could push price beyond $2 for Alaska dive fleet". National Fisherman. http://www.nationalfisherman.com/2008.asp?ItemID=1800&pcid=373&cid=375&archive=yes. Retrieved 2008-08-01.  

Wikispecies

Up to date as of January 23, 2010

From Wikispecies

Taxonavigation

Main Page
Cladus: Eukaryota
Supergroup: Unikonta
Cladus: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Echinodermata
Subphylum: Eleutherozoa
Superclassis: Cryptosyringida
Classis: Holothuroidea
Subclassis: Apodacea - Aspidochirotacea - Dendrochirotacea

References

  • Kerr, A. 2000: Evolution and Systematics of Holothuroidea (Echinodermata). Thesis, Yale University.
  • O’Loughlin, P.M. 2009: BANZARE holothuroids (Echinodermata: Holothuroidea). Zootaxa, 2196: 1-18. Abstract & excerpt PDF
  • Smiley, S. 1994: Holothuroidea. In: Harrison, F. W. and F.-S. Chia, eds. Microscopic Anatomy of Invertebrates. Volume 14. Echinodermata. New York: Wiley-Liss, 401-471.
  • Smith, A.B. 1997: Echinoderm larvae and phylogeny. Annual Review of Ecology and Systematics, 28: 219-241.

Vernacular names

Česky: Sumýši
Deutsch: Seegurken
Ελληνικά: Ολοθουροειδή (Αγγούρια της θάλασσας)
English: Sea cucumbers
Español: Pepinos de mar
Italiano: Cetrioli di mare, Oloturie
Slovenčina: Holotúrie
Suomi: Merimakkarat
Wikimedia Commons For more multimedia, look at Holothuroidea on Wikimedia Commons.

Simple English

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