Tunicate: Difference between revisions

From Wikipedia, the free encyclopedia
Browse history interactively
← Previous editNext edit →
Content deleted Content added
VisualWikitext
→‎Classification: New information and reference
Replacing automatic taxobox with manual one
Line 1: Line 1:
{{Taxobox
{{Automatic taxobox
| fossil_range={{fossil range|520|0|earliest=555|text=Cambrian "Stage 3"–recent |ref=<ref name="Fedonkin2012">{{cite doi|10.1134/S0031030112010042}}</ref>}}
⚫
| taxon = Tunicata
| name = Tunicate
| oldest fossil = Cambrian stage 3
| image = Sea-tulip.jpg
| image = Sea-tulip.jpg
| image_width = 250px
| image_caption = Sea tulips ''[[Pyura spinifera]]''
| image_caption = Sea tulips ''[[Pyura spinifera]]''
| regnum = [[Animal]]ia
⚫
| authority = [[Jean-Baptiste Lamarck|Lemarck]] 1816<ref name=WoRMS>{{cite WoRMS |author=Sanamyan, Karen |year=2013 |title=Tunicata |id=146420 |accessdate=2013-04-04 |db=}}</ref>
| phylum = [[Chordata]]
| display children = 1
⚫
| subphylum = '''Tunicata'''
⚫
| subphylum_authority = [[Jean-Baptiste Lamarck|Lemarck]] 1816<ref name=WoRMS>{{cite WoRMS |author=Sanamyan, Karen |year=2013 |title=Tunicata |id=146420 |accessdate=2013-04-04 }}</ref>
| subdivision_ranks = [[Class (biology)|Classes]]<ref name=WoRMS/><ref name="Tatian"/>
| subdivision =
*[[Enterogona]]
*[[Ascidiacea]]
*[[Thaliacea]]
}}
}}


Revision as of 09:52, 9 April 2013

Tunicate
Temporal range: Cambrian "Stage 3"–recent[1]
Sea tulips Pyura spinifera
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Subphylum:
Tunicata

Classes[2][3]

Tunicates, previously known as Urochordata or urochordates, are members of the Tunicata, a subphylum of the phylum Chordata. They are marine filter feeders with a sac-like body structure. In their respiration and feeding they take in water through an incurrent (or inhalant) siphon and expel the filtered water through an excurrent (or exhalant) siphon. Most adult tunicates are sessile and attached to rocks or similarly suitable surfaces on the ocean floor; others such as salps, doliolids and pyrosomes swim in the pelagic zone as adults. Various species are commonly known as sea squirts, sea pork or sea tulips.

The Tunicata appear to have evolved in the early Cambrian period. Despite their simple appearance and very different adult form, they are a sister subphylum to the Vertebrata.

Body form

There are about 2,000 species of tunicate in the world's oceans, with fewer than 100 being found at depths greater than 200 metres (660 ft). Some are solitary animals leading a sessile existence attached to the seabed, but others are colonial and a few are pelagic. Some are supported by a stalk but most are attached directly to the substrate, which may be a rock, shell, coral, seaweed, mangrove root, dock, piling or ship's hull. They come in a range of solid or translucent colours and may resemble seeds, grapes, peaches, barrels or bottles. One of the largest is Pyura pachydermatina, a sea tulip, which can be over a metre (yard) tall.[4]

Colonies come in a range of forms and degrees of integration. In the simplest systems, the zooids are widely separated but linked together by a stolon which grows across the seabed. Other species have the zooids growing closeer together in a tuft or clustered together and sharing a common base. The most advanced colonies involve the integration of the zooids into a common structure surrounded by the tunic. These may have separate buccal siphons and a single central atrial siphon and may be organized into larger systems with hundreds of star-shaped units. Often the zooids in a colony are tiny, but very numerous, and the colonies can form large encrusting or mat-like patches.[4]

Body structure

Diagrammatic section through an ascidian:
a) buccal siphon; b) atrial siphon; c) tunic d) mantle; e) pharynx; f) gullet; g) stomach; h) anus; i) endostyle
Oikopleura cophocerca in its "house"; arrows indicate water movement and (x) the lateral, reticulated parts of the "house"

By far the largest class of tunicates is the Ascidiacea. The body of an ascidiacean is surrounded by a test or tunic, from which the class derives its name. This varies in thickness between species but may be tough, resembling cartilage, thin and delicate or transparent and gelatinous. The tunic is composed of proteins and complex carbohydrates and includes tunicin, a variety of cellulose. The tunic is unique among invertebrate exoskeletons in that it can grow as the animal enlarges and does not need to be periodically shed. Inside the tunic is the body wall or mantle. This is composed of connective tissue, muscle fibres, blood vessels and nerves. There are two openings in the body wall, the buccal siphon at the top through which water flows into the interior and the atrial siphon on the ventral side through which it is expelled. A large pharynx occupies most of the interior of the body. It is lined by a pharyngeal net with a ciliated groove known as an endostyle on its ventral surface. This is connected via the gullet to a loop of gut which terminates near the atrial siphon, from where faecal matter is swept away with the exhalent water. The walls of the pharynx are perforated by several bands of pharyngeal slits through which water escapes into the surrounding water-filled cavity, the atrium. This is criss-crossed by various mesenteries which provide support for the pharynx, preventing it from collapsing, and also support the other organs.[4]

Thaliacea, the other main class of tunicates, is characterised by free-swimming, pelagic individuals. They are all filter feeders using a pharyngeal mucous net to catch their prey. The pyrosomes are bioluminous colonial tunicates with a hollow cylindrical structure. The buccal siphons are on the outside and the atrial siphons inside. There are about ten species, and all are found in the tropics. The twenty three species of doliolids are small, mostly under 2 centimetres (0.8 in)* long. They are solitary, have the two siphons at opposite ends of their barrel-shaped bodies and swim by jet propulsion. The forty species of salps are also small, under 4 centimetres (1.6 in)* long, and found in the surface waters of both warm and cold seas. They also move by jet propulsion and often form long chains by budding off new individuals.[4]

A third class, Larvacea (or Appendicularia), is the only group of tunicates to retain its chordate characteristics in the adult state. There are seventy species of larvacean which superficially resemble the tadpole larvae of amphibians, although the tail is at right angles to the body. The notochord is retained and the animals, mostly under a centimetre long, are propelled by undulations of the tail. They secrete an external mucous net known as a house which may completely surround them and is very efficient at trapping planktonic particles.[4]

Physiology and internal anatomy

Like other chordates, tunicates have a notochord during their early development, but by the time they have completed their larval stages they have lost all myomeric segmentation throughout the body. As members of the Chordata they are true Coelomata with endoderm, ectoderm and mesoderm, but they do not develop very clear coelomic body-cavities if any at all. Whether they do or not, by the end of their larval development all that remain are the pericardial, renal, and gonadal cavities of the adults. Except for the heart, gonads, and pharynx (or branchial sac), the organs are enclosed in a membrane called an epicardium, which is surrounded by the jelly-like mesenchyme. Tunicates begin life in a mobile larval stage that resembles a tadpole. A minority of species, those in the Larvacea, retain the general larval form throughout life, but most Tunicata very rapidly settle down and attach themselves to a suitable surface, later developing into a barrel-like and usually sedentary adult form. The Thaliacea however, are pelagic throughout their lives and may have complex life cycles.[5]

Tunicates have a well-developed heart and circulatory system. The heart is a double U-shaped tube situated just below the gut. The blood vessels are simple connective tissue tubes and there are several types of corpuscle. The blood may appear pale green but this is not due to any respiratory pigments and oxygen is transported dissolved in the plasma. Exact details of the circulatory system are unclear but the gut, pharynx, gills, gonads and nervous system seem to be arranged in series rather than in parallel as happens in most other animals. Every few minutes the heart stops beating and then restarts, pumping fluid in the reverse direction.[4]

Tunicate blood has some unusual features. In some species of Ascidiidae and Perophoridae it contains high concentrations of the transitional metal vanadium and vanadium-associated proteins in vacuoles in blood cells known as vanadocytes. Some tunicates can concentrate vanadium up to a level ten million times that of the surrounding seawater. It is stored in a +3 oxidation form that requires a pH of less than 2 for stability and this is achieved by the vacuoles also containing sulphuric acid. The vanadocytes are later deposited just below the outer surface of the tunic where it is thought that their presence deters predation. Other species of tunicate concentrate lithium, iron, niobium and tantalum which may serve a similar function.[4]

Tunicates lack the kidney-like metanephridial organs typical of deuterostomes. Most have no excretory structures but rely on the diffusion of ammonia across their tissues to rid themselves of nitrogenous waste but some have a simple excretory system. The typical renal organ is a mass of large clear-walled vesicles that occupy the rectal loop, and the structure has no duct. Each vesicle is a remnant of a part of the primitive coelom, and its cells extract nitrogenous waste matter from circulating blood. They accumulate the wastes inside the vesicles as urate crystals and do not have any obvious means of disposing of the material during their lifetime.[5]

Tunicates are unusual among animals in that they produce a large fraction of their tunic and some other structures in the form of cellulose. The production in animals of cellulose is so unusual that at first some researchers denied its presence outside plants, but for some time now it has been accepted that it occurs in the dermis of mammals. However, the tunicates are unique in their scale of application and production of the material, and possibly in having a functional cellulose synthesising enzyme.[6] When in 1845 Carl Schmidt first announced the presence in the test of some Ascidians of a substance very similar to cellulose, he called it "tunicine", but it now is recognised as cellulose rather than any alternative substance.[7][8][9]

Feeding

Most tunicates are suspension feeders, capturing suspended particles and plankton by filtering sea water through their siphons via their pharyngeal slits.

Tunicates have two openings in their body cavity: an incurrent and an excurrent siphon. The incurrent siphon takes in water plus any food it contains, and the excurrent siphon expels wastes plus sieved water. The primary food source for most tunicates is plankton that gets entangled in mucus secreted from the endostyle. The tunicate's pharynx, or branchial sac, is lined with ciliated epithelium. The action of the cilia passes food particles and plankton down to the esophagus and also passes the stream of water from the inhalant to the exhalant siphon. The gut is U-shaped and also ciliated, and the anus opens into the dorsal or cloacal part of the peribranchial cavity near the atrial aperture, where the constant stream of water carries wastes to the exterior.

Some ascidians that live on soft sediments are detritivores. A few deep water species such as Megalodicopia hians are sit-and-wait predators, trapping tiny crustacea, nematodes and other small invertebrates with the muscular lobes which surround their buccal siphons. Certain tropical species in the family Didemnidae have symbiotic green algae or cyanobacteria in their tunics and one of these symbionts, Prochloron, is unique to tunicates. It is assumed that excess photosynthetic products are available to the host.[4]

Life cycle

Comparison of frog tadpole (above)
and tunicate tadpole (below)

Tunicates are hermaphrodites and each has a single ovary and testis. In some solitary species, sperm and eggs are shed into the sea and the larvae are planktonic. In others, especially in colonial species, sperm is released into the water and is drawn into the atria of other individuals with the incoming water current. Fertilisation takes place here and the eggs are brooded through their early developmental stages.[5] Some larval forms appear very much like primitive chordates with a notochord (stiffening rod). Superficially, the larva resembles a small tadpole. It swims with a tail and may have a simple eye, or ocellus, and balancing organ, or statolith.[10]

The larval form ends when the tunicate finds a suitable rock to affix to and cements itself in place. The larval form is not capable of feeding, though it may have a rudimentary digestive system,[10] and is only a dispersal mechanism. Many physical changes occur to the tunicate's body during metamorphosis, one of the most interesting being the digestion of the cerebral ganglion, which controls movement and is the equivalent of the human brain. From this comes the common saying that the sea squirt "eats its own brain".[11] In some classes, the adults remain pelagic (swimming or drifting in the open sea), although their larvae undergo similar metamorphoses to a higher or lower degree.[5]

During embryonic development, tunicates exhibit determinate cleavage, where the fate of the cells is set early on with reduced cell numbers and genomes that are rapidly evolving. In contrast, the amphioxus and vertebrates show cell determination relatively late in development and cell cleavage is indeterminate. The genome evolution of amphioxus and vertebrates is also relatively slow.[12]

Colonial forms also increase the size of the colony by budding off new individuals to share the same tunic.[13]

Taxonomy

Clavelina moluccensis, the bluebell tunicate
Tunicate colonies of Botrylloides violaceus with oral tentacles at openings of oral siphons visible
Colonial tunicate with multiple openings in a single tunic

Urochordata is a junior synonym of Tunicata which was established by Lamarck in 1816. Balfour introduced the name Urochorda in 1881 in order to emphasize the affinity of the group to other chordates,.[14] No doubt largely because of his influence, various authors supported the term, either as such, or as "Urochordata", but the usage is invalid because "Tunicata" has precedence and there never were grounds for superseding the name. Accordingly the current (formally correct) trend is to abandon the name Urochorda or Urochordata in favour of the original Tunicata, and the name Tunicata is almost invariably used in modern scientific works. It is accepted as valid by the World Register of Marine Species[15] and by ITIS, the Integrated Taxonomic Information System.[16]

Various species are commonly known as sea tulips, sea squirts[17] or sea pork.[18]

Classification

Tunicates are more closely related to craniates (including hagfish, lampreys, and jawed vertebrates) than to lancelets, echinoderms, hemichordates, Xenoturbella or other invertebrates.[19][20][21]

The clade comprising Tunicates and Vertebrates is called Olfactores.[22]

The Tunicata contains about 3,000 species, traditionally divided into the following classes:

Members of the Sorberacea were included in Ascidiacea in 2011 as a result of rDNA sequencing studies.[3]

Although the traditional classification is provisionally accepted, newer evidence suggests that the Ascidiacea is an artificial group of paraphyletic status.[23][24]

Sea squirts have been involved in a controversy about the extent to which cross-species gene transfer and hybridization have influenced animal evolution. In 1990, Donald I. Williamson of the University of Liverpool (U.K.) fertilised sea squirt (Ascidia mentula) eggs with sea urchin (Echinus esculentus) sperm resulting in fertile adults that resembled urchins,[25] but Michael W. Hart of Simon Fraser University failed to find sea-squirt DNA in tissue samples from the supposed hybrids.[26] Williamson claims to have repeated the experiment with sea urchin eggs and sea squirt sperm, producing sea urchin larvae which developed into squirt-like juveniles.[27] These findings have been disputed. A multi-taxon molecular study in 2010 suggested that sea squirts are descended from a hybrid between a chordate and a likely extinct protostome ancestor which took place at a time before the diversification of nematodes and arthropods. This study also examined whether there was any evidence of a sea urchin and tunicate hybridization event that could possibly explain the distribution of genes in modern sea squirts. None could be found.[28][29]

Fossil record

The star-shaped holes (Catellocaula vallata) in this Upper Ordovician bryozoan represent a tunicate preserved by bioimmuration in the bryozoan skeleton.

Undisputed fossils of tunicates are rare. The best known (and earliest) is Shankouclava shankouense from the Lower Cambrian Maotianshan Shale at Shankou village, Anning, near Kunming (South China).[30] There is also a common bioimmuration of a tunicate (Catellocaula vallata) found in Upper Ordovician bryozoan skeletons of the upper midwestern United States. [31]

There are also two enigmatic species from the Ediacaran period - Ausia fenestrata from the Nama Group of Namibia and a second new Ausia-like genus from the Onega Peninsula, White Sea of northern Russia. Results of new study have shown possible affinity of these Ediacaran organisms to the ascidians.[32][33] These two organisms lived in the shallow waters of a sea, slightly more than 555-548 million years ago and are believed to be the oldest evidence of the chordate lineage of metazoans.[33]

A Precambrian fossil known as Yarnemia is thought to be a tunicate however this has been disputed. Fossils of tunicates are rare because their bodies decay soon after death but in some tunicate families, microscopic spicules are present which may be preserved as microfossils. These spicules have occasionally been found in Jurassic and later rocks but as few palaeontologists are familiar with them, they may have been mistaken for sponge spicules.[34]

Invasive species

Over the past few decades, tunicates (notably of the genera Didemnum and Styela) have been invading coastal waters in many countries. The carpet tunicate (Didemnum vexillum) has taken over a 6.5 square miles (17 km2) area of the seabed on the Georges Bank off the north east coast of North America, covering stones, molluscs and other stationary objects in a dense mat.[35] D. vexillum, Styela clava and Ciona savignyi have appeared and are thriving in Puget Sound and Hood Canal in the Pacific Northwest.[36]

Introduction of invasive tunicates usually occurs as fouling organisms on the hulls of ships and possibly the larvae have arrived in ballast water. Another possible route is on the shells of molluscs brought in for aquaculture.[36] Current research indicates that many tunicates previously thought to be indigenous to Europe and the Americas are, in fact, invaders. Some of these invasions may have occurred centuries or even millennia ago. In some areas, tunicates are proving to be a major threat to aquaculture operations.[37]

Medical uses

Tunicates contain a host of potentially useful chemical compounds, including:

In the May 2007 issue of the FASEB Journal, researchers from Stanford University showed that tunicates can correct abnormalities over a series of generations, and they suggest that a similar regenerative process may be possible for humans. The mechanisms underlying the phenomenon may lead to insights about the potential of cells and tissues to be reprogrammed and regenerate compromised human organs. Gerald Weissman, editor-in-chief of the journal, said "This study is a landmark in regenerative medicine; the Stanford group has accomplished the biological equivalent of turning a sow's ear into a silk purse and back again."[38]

As food

Sea squirts for sale at a market, Busan, South Korea
Main article: Ascidiacea § Culinary

Various Ascidiacea species are consumed as food around the world. In Japan and Korea, the sea pineapple (Halocynthia roretzi) is the main species eaten. There is an aquaculture industry where it is cultivated on dangling cords made of palm fronds. In 1994, over 42,000 tons were produced but since then there have been problems with mass mortality events and only 4,500 tons were produced in 2004.[39]

Other uses

The use of tunicates as a source of biofuel is being researched. The cellulose body wall can be broken down and converted into ethanol and other parts of the animal are protein-rich and can be converted into fish feed. The researchers believe that culturing tunicates on a large scale is possible and that the economics of doing so are attractive. They believe that, as tunicates have few predators, their removal from the sea may not have profound ecological impacts. Being sea-based, their production does not compete with food production as does the cultivation of land-based crops for biofuel projects.[40]

Some tunicates are used as model organisms. The species Ciona intestinalis and Ciona savignyi have been used for developmental studies. Both species' mitochondrial[41][42] and nuclear[43][44] genomes have been sequenced. The nuclear genome of the appendicularian Oikopleura dioica appears to be one of the smallest among metazoans[45] and this species has been used to study gene regulation and the evolution and development of chordates.[46]

References

  1. ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1134/S0031030112010042, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1134/S0031030112010042 instead.
  2. ^ a b Sanamyan, Karen (2013). "Tunicata". WoRMS. World Register of Marine Species. Retrieved 4 April 2013.
  3. ^ a b Tatián, Marcos; Lagger, Cristian; Demarchi, Milagros; Mattoni, Camilo (2011). "Molecular phylogeny endorses the relationship between carnivorous and filter-feeding tunicates (Tunicata, Ascidiacea)". Zoologica Scripta. 40 (6): 603–612. doi:10.1111/j.1463-6409.2011.00493.x.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  4. ^ a b c d e f g h Ruppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004). Invertebrate Zoology, 7th edition. Cengage Learning. p. 947. ISBN 81-315-0104-3.{{cite book}}: CS1 maint: multiple names: authors list (link)
  5. ^ a b c d Dorit, R. L.; Walker, W. F.; Barnes, R. D. (1991). Zoology. Saunders College Publishing. pp. 802–804. ISBN 978-0-03-030504-7. Cite error: The named reference "Dorit" was defined multiple times with different content (see the help page).
  6. ^ Matthysse, Ann G.; Deschet, Karine; Williams, Melanie; Marry, Mazz; White, Alan R.; Smith, William C. (2004). "A functional cellulose synthase from ascidian epidermis". PNAS. 101 (4): 986–991. doi:10.1073/pnas.0303623101.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  7. ^ Hirose, Euichi; Nakashima, Keisuke; Nishino, Atsuo (2011). "Is there intracellular cellulose in the appendicularian tail epidermis? A tale of the adult tail of an invertebrate chordate". Communicative & Integrative Biology. 4 (6): 768–771.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  8. ^ Hall, D. A.; Saxl, Hedwig (1961). "Studies of Human and Tunicate Cellulose and of their Relation to Reticulin". Proceedings of the Royal Society of London. 155 (959): 202–217. doi:10.1098/rspb.1961.0066}.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ Sasakura, Yasunori; Nakashima, Keisuke; Awazu, Satoko; Matsuoka, Terumi; Nakayama, Akie; Azuma, Jun-ichi; Satoh, Nori (2005). "Transposon-mediated insertional mutagenesis revealed the functions of animal cellulose synthase in the ascidian Ciona intestinalis". 102 (42): 15134–15139. doi:10.1073/pnas.0503640102. {{cite journal}}: Cite journal requires |journal= (help)CS1 maint: multiple names: authors list (link)
  10. ^ a b Cavanihac, Jean-Marie (2000). "Tunicates extraordinaire". Microscope UK. Retrieved 7 December 2011.
  11. ^ "Brainless Fish in Topless Bar". Fast Company. 30 April 1999. Retrieved 7 December 2011.
  12. ^ Holland, Linda Z. (2007). "Developmental biology: A chordate with a difference". Nature. 447 (1): 153–155. doi:10.1038/447153a;.{{cite journal}}: CS1 maint: extra punctuation (link)
  13. ^ Parmentier, Jan (1998). "Botryllus: A colonial ascidian". Microscope UK. Retrieved 7 April 2013.
  14. ^ Foster, M. (ed.); Sedgwick, Adam (ed.); The Works of Francis Maitland Balfour. Vol. III. Memorial edition. Pub: Macmillan and co. 1885. May be downloaded from [1]
  15. ^ Tunicata World Register of Marine Species. Retrieved 2011-11-12.
  16. ^ Tunicata Lamarck, 1816 Integrated Taxonomic Information System. Retrieved 2011-11-12.
  17. ^ Branch, G. M.; Griffiths, C. L.; Branch, M. L.; Beckley, L. E. (2010). Two oceans: a guide to the marine life of Southern Africa. Cape Town: Struik Nature. ISBN 978-1-77007-772-0.{{cite book}}: CS1 maint: multiple names: authors list (link)
  18. ^ "Gulf Specimen Marine Laboratory: Sea Squirts and Sea Pork". Retrieved 5 April 2013.
  19. ^ Delsuc, F.; Brinkmann, H.; Chourrout, D.; Philippe, H. (2006). "Tunicates and not cephalochordates are the closest living relatives of vertebrates". Nature. 439 (7079): 965–968. doi:10.1038/nature04336. PMID 16495997.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  20. ^ Delsuc, F.; Tsagkogeorga, G.; Lartillot, N.; Philippe, H. (2008). "Additional molecular support for the new chordate phylogeny". Genesis. 46 (11): 592–604. doi:10.1002/dvg.20450. PMID 19003928.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  21. ^ Singh, T. R.; Tsagkogeorga, G.; Delsuc, F.; Blanquart, S.; Shenkar, N.; Loya, Y.; Douzery, E. J.; Huchon, D. (2009). "Tunicate mitogenomics and phylogenetics: peculiarities of the Herdmania momus mitochondrial genome and support for the new chordate phylogeny". BMC Genomics. 10: 534. doi:10.1186/1471-2164-10-534. PMC 2785839. PMID 19922605.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  22. ^ Jefferies, R. P. S. (1991) in Biological Asymmetry and Handedness (eds Bock, G. R.; Marsh, J.) pp. 94-127 (Wiley, Chichester).
  23. ^ Zeng, L.; Swalla, B. J. (2005). "Molecular phylogeny of the protochordates: chordate evolution". Can. J. Zool. 83: 24–33. doi:10.1139/z05-010.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  24. ^ Tsagkogeorga, G.; Turon, X.; Hopcroft, R. R.; Tilak, M. K.; Feldstein, T.; Shenkar, N.; Loya, Y.; Huchon, D.; Douzery, E. J.; Delsuc, F. (2009). "An updated 18S rRNA phylogeny of tunicates based on mixture and secondary structure models". BMC Evolutionary Biology. 9: 187. doi:10.1186/1471-2148-9-187. PMC 2739199. PMID 19656395.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  25. ^ Williamson, D.I.; Vickers, S.E. (2007). "The Origins of Larvae: Differences in the forms of larvae and adults may reflect fused genomes" (PDF). American Scientist. 95 (6): 509–517.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  26. ^ Hart, M.W. (1996). "Testing cold fusion of phyla: Maternity in a tunicate × sea urchin hybrid determined from DNA comparisons". Evolution. 50 (4): 1713–1718.
  27. ^ Williamson, D.I. (in press). Larval transfer: experimental hybrids. In: Margulis, L. and Asikainen, C.A. (editors), Chimeras and Consciousness: Evolution of Sensory Systems. White River Junction, Vermont: Chelsea Green Publishing Co. (Cit. in: Williamson, D. I.; Vickers, S. E. Larval Transfer: A recent evolutionary theory. Ms. submitted to American Scientist.)
  28. ^ Syvanen, M.; Ducore, J. (2010). "Whole genome comparisons reveals a possible chimeric origin for a major metazoan assemblage". Journal of Biological Systems. 18: 261–275. doi:10.1142/S0218339010003408.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  29. ^ Lawton, G. (21 January 2009). "Uprooting Darwin's tree" (PDF). New Scientist. Retrieved 19 April 2013.
  30. ^ Chen, Jun-Yuan; Huang, Di-Ying; Peng, Qing-Qing; Chi, Hui-Mei; Wang,Xiu-Qiang; Feng, Man (2003). "The first tunicate from the Early Cambrian of South China". Proceedings of the National Academy of Sciences. 100 (14): 8314–8318. doi:10.1073/pnas.1431177100. PMC 166226. PMID 12835415.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  31. ^ Palmer, T. J.; Wilson, M. A. (1988). "Parasitism of Ordovician bryozoans and the origin of pseudoborings" (PDF). Palaeontology. 31: 939–949.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  32. ^ Vickers-Rich P. (2007). "Chapter 4. The Nama Fauna of Southern Africa". In: Fedonkin, M. A.; Gehling, J. G.; Grey, K.; Narbonne, G. M.; Vickers-Rich, P. "The Rise of Animals: Evolution and Diversification of the Kingdom Animalia", Johns Hopkins University Press. pp. 69-87
  33. ^ a b Fedonkin, M. A.; Vickers-Rich, P.; Swalla, B.; Trusler, P.; Hall, M. (2008). "A Neoproterozoic chordate with possible affinity to the ascidians: New fossil evidence from the Vendian of the White Sea, Russia and its evolutionary and ecological implications". HPF-07 Rise and fall of the Ediacaran (Vendian) biota. International Geological Congress - Oslo 2008.
  34. ^ "Introduction to the Urochordata". University of California Museum of Paleontology. Retrieved 7 April 2013.
  35. ^ "Have You Seen This Tunicate?". NOAA Fisheries Service. 19 November 2004. Retrieved 7 December 2011.
  36. ^ a b Dornfeld, Ann (1 May 2008). "Invasive Tunicates of Washington State". NPR. Retrieved 6 April 2013.
  37. ^ "Marine Nuisance Species". Woodshole Science Center. Retrieved 7 December 2011.
  38. ^ "Sea Squirt, Heal Thyself: Scientists Make Major Breakthrough in Regenerative Medicine". Sciencedaily.com. 24 April 2007. Retrieved 7 December 2011.
  39. ^ "Sea squirt". Korea-US Aquaculture. Retrieved 6 April 2013.
  40. ^ "Biofuel made from marine filter feeders? Tunicates usable as source of biofuels". Cleantechnica. 26 March 2013. Retrieved 6 April 2013.
  41. ^ Iannelli, F.; Pesole, G.,; Sordino, P.; Gissi, C. (2007). "Mitogenomics reveals two cryptic species in Ciona intestinalis". Trends Genet. 23 (9): 419–422. doi:10.1016/j.tig.2007.07.001. PMID 17640763.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  42. ^ Yokobori, S.; Watanabe, Y.; Oshima, T. (2003). "Mitochondrial genome of Ciona savignyi (Urochordata, Ascidiacea, Enterogona): Comparison of gene arrangement and tRNA genes with Halocynthia roretzi mitochondrial genome". J. Mol. Evol. 57 (5): 574–587. doi:10.1007/s00239-003-2511-9. PMID 14738316. {{cite journal}}: Cite has empty unknown parameter: |month= (help)CS1 maint: multiple names: authors list (link)
  43. ^ Dehal, P.; Satou, Y.; Campbell, R. K.; Chapman, J., Degnan, B., De Tomaso, A.; Davidson, B.; Di Gregorio, A.; Gelpke, M.; Goodstein, D. M.; Harafuji, N.; Hastings, K. E.; Ho, I.; Hotta, K.; Huang, W.; Kawashima, T.; Lemaire, P.; Martinez, D.; Meinertzhagen, I. A.; Necula, S.; Nonaka, M.; Putnam, N.; Rash, S.; Saiga, H.; Satake, M.; Terry, A.; Yamada L.; Wang, H. G.; Awazu, S.; Azumi, K.; Boore, J.; Branno, M.; Chin-Bow, S.; DeSantis, R.; Doyle, S., Francino, P.; Keys, D. N.; Haga, S.; Hayashi, H.; Hino, K.; Imai, K. S.; Inaba, K.; Kano, S.; Kobayashi, K.; Kobayashi, M.; Lee, B. I.; Makabe, K. W.; Manohar, C.; Matassi, G.; Medina, M.; Mochizuki, Y.; Mount, S.; Morishita, T.; Miura, S.; Nakayama, A.; Nishizaka, S.; Nomoto, H.; Ohta, F.; Oishi, K.; Rigoutsos, I.; Sano, M.; Sasaki, A.; Sasakura, Y.; Shoguchi, E.; Shin-i, T.; Spagnuolo, A.; Stainier, D.; Suzuki, M. M.; Tassy, O.; Takatori, N.; Tokuoka, M.; Yagi, K.; Yoshizaki, F.; Wada, S.; Zhang C.; Hyatt, P. D.; Larimer, F.; Detter, C.; Doggett, N.; Glavina, T.; Hawkins, T.; Richardson, P.; Lucas, S.; Kohara, Y.; Levine, M.; Satoh, N.; Rokhsar, D. S. (2002). "The draft genome of Ciona intestinalis: insights into chordate and vertebrate origins". Science. 298 (5601): 2157–2167. doi:10.1126/science.1080049. PMID 12481130.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  44. ^ Small, K. S.; Brudno, M.; Hill, M. M.; Sidow, A. (2007). "A haplome alignment and reference sequence of the highly polymorphic Ciona savignyi genome". Genome Biol. 8 (3): R41. doi:10.1186/gb-2007-8-3-r41. PMC 1868934. PMID 17374142. {{cite journal}}: Cite has empty unknown parameter: |month= (help)CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  45. ^ Seo, H. C.; Kube, M.; Edvardsen, R. B.; Jensen, M. F.; Beck, A.; Spriet, E.; Gorsky, G.; Thompson. E. M.; Lehrach, H.; Reinhardt, R.; Chourrout, D. (2001). "Miniature genome in the marine chordate Oikopleura dioica". Science. 294 (5551): 2506–2506. doi:10.1126/science.294.5551.2506. PMID 11752568.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  46. ^ Clarke, T.; Bouquet, JM; Fu, X; Kallesøe, T.; Schmid, M; Thompson, E.M. (2007). "Rapidly evolving lamins in a chordate, Oikopleura dioica, with unusual nuclear architecture". Gene. 396 (1): 159–169. doi:10.1016/j.gene.2007.03.006. PMID 17449201.

External links

Wikimedia Commons has media related to Tunicata.
Wikispecies has information related to Urochordata.

Retrieved from "https://en.wikipedia.org/w/index.php?title=Tunicate&oldid=549484215"