tunicate


tunicate
/tooh"ni kit, -kayt', tyooh"-/, n.
1. Zool. any sessile marine chordate of the subphylum Tunicata (Urochordata), having a saclike body enclosed in a thick membrane or tunic and two openings or siphons for the ingress and egress of water.
adj. Also, tunicated.
2. (esp. of the Tunicata) having a tunic or covering.
3. of or pertaining to the tunicates.
4. Bot. having or consisting of a series of concentric layers, as a bulb.
[1615-25; < L tunicatus wearing a tunic. See TUNIC, -ATE1]

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Any of some 2,000 species (chordate subphylum Tunicata, or Urochordata) of small marine invertebrates that are abundant worldwide.

Tunicates are either sessile (permanently attached), free-swimming, or planktonic (floating). The name tunicate derives from a secreted protective covering (the tunic) containing cellulose. Floating species often form colonies that may be 13 ft (4 m) long. Some free-swimming species are too small to see. Adults filter feed on microorganisms. Sessile forms growing on ships' hulls may be a nuisance, but some species are pharmaceutically useful. See also sea squirt.

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▪ chordate subphylum
Introduction
also called  urochordate,  

      any member of the subphylum Tunicata (Urochordata) of the phylum Chordata. Small marine animals, they are found in great numbers throughout the seas of the world.

      Adult members are commonly embedded in a tough secreted tunic containing cellulose (a glucose polysaccharide not normally found in animals). The less modified forms are benthic (bottom-dwelling and sessile), while the more advanced forms are pelagic (floating and swimming in open water). A characteristic tadpole larva develops in the life cycle, and in one group (the appendicularians, or larvaceans) the adult closely resembles this larva, which has many features in common with other chordates. Most chordate features disappear at metamorphosis.

General features

Size range and diversity of structure
      The tunicates are divided into three classes: Ascidiacea (ascidians, or sea squirts (sea squirt)), Appendicularia (Larvacea (larvacean)), and Thaliacea. Ascidians are largely benthic animals. They often form colonies, comprising a few to many individuals (zooids), which reach up to two metres in length. Solitary (noncolonial) forms range from one millimetre to over 20 centimetres in length. The adult appendicularian resembles the tadpole larva of other tunicates. The body is enveloped in a “house,” with which the animal nets food. Small (usually around five millimetres in length, including the tail) and simple, appendicularians do not form colonies. They spend their entire lives in the open sea. The thaliaceans (pyrosomes, dolioloids, and salps (salp)) are also pelagic. Their structure suggests that they are ascidians modified in adaptation to conditions in open water. They have specialized modes of reproduction, sometimes with a complicated alteration of sexual and asexual phases. Pyrosomes form long, tubular colonies. Dolioloids and salps occur both as solitary individuals and as chains.

Distribution and abundance
      Tunicates are distributed in ocean waters from the polar regions to the tropics. Free-swimming tunicates are found throughout the oceans as plankton, while sessile forms grow mainly on solid surfaces such as wharf piles, ship hulls, rocks, and the shells of various sea creatures.

Importance
      Although rarely eaten by humans, tunicates are an important link in the food chain and thus indirectly provide humans with a source of food. Tunicates contain some unusual chemicals, and some of these may prove useful as drugs. Some tunicates are fouling organisms that grow on ships' hulls. Their main interest to humans is in providing clues to the possible ancestry of vertebrates.

Natural history

Reproduction and life cycle
      With rare exceptions, tunicates are hermaphrodites, but reproduction may be by sexual or asexual (budding) means. In general, hermaphroditic animals do not self-fertilize (i.e., provide both the male and female gametes) if they can avoid doing so, a rule that seems also to be true of tunicates. In primitive forms the eggs are fertilized, and development takes place, in the surrounding water, but often embryos are retained in the female's atrium or elsewhere until the larva is developed.

      The larval stage is brief; the larva does not feed, but concentrates on finding an appropriate place for the adult to live. In keeping with this motile phase, the muscular tail comprises two-thirds of the larval body; it is supported by a notochord and contains a nerve cord. Gravity- and light-sensitive sensory vesicles along the dorsal surface of the larval body orient the animal as it swims. After a period of up to a few days, the larva will settle and attach itself to a surface using three anterior adhesive papillae. As the larva metamorphoses into an adult, the tail resorbs, providing food reserves for the developing animal. Free-swimming tunicates metamorphose without attachment.

      Colonies are formed by asexual reproduction, with zooids usually being formed by budding. In thaliaceans, two groups (dolioloids and salps) have a complex system of alternating phases; the first phase reproduces by budding, and the resulting individuals may release sperm and eggs.

Locomotion
      Tadpole larvae and appendicularians swim by undulating the tail, which contains a stiff notochord. Despite their sessile life-styles, some adult ascidians can move by attaching with one area of the body and letting go with another. Movement of colonies up to 1.5 centimetres per day has been recorded. In thaliaceans an exhalant current of water, which in dolioloids and salps is combined with a strong muscular contraction, creates a jet stream that propels the animal forward.

Food and feeding
      An internal mucous sheet, secreted by the endostyle, allows ascidians and thaliaceans to utilize a variety of organisms, especially small plants (phytoplankton) as their food source (see below Internal features: Digestion, nutrition, and excretion). Some trap small animals. The feeding mechanism is different in appendicularians. Glands on the surface of the body secrete a complex house made up of mucus, which surrounds the animal. Undulations of the tail produce a feeding current that draws water into the house and through a fine sheet of mucus, which serves as a net to filter the food. Appendicularians feed on microscopic organisms (nannoplankton).

Associations
      Tunicates often host various parasitic animals. Some tunicates, especially in the tropics, live symbiotically with unicellular plants and blue-green algae that may supply them with food.

Form and function

General features
      A tunicate tadpole larva contains several chordate features, such as the notochord, dorsal nerve cord, and tail. These features are lost, however, as the larva metamorphoses into the adult form. The tunicate larva has special organs of sense and attachment, which it uses to find and occupy a suitable habitat. Once the larva has attached to a substrate by its anterior end, the larval features quickly regress and considerable changes in size and proportion of parts take place. For example, the notochord, nerve cord, and most of the tail are generally resorbed within one day. The area between the mouth and the point of attachment grows rapidly until the mouth comes to be directed away from the point of attachment, which now becomes the posterior end of the animal. The atrium usually forms from a pair of pouches that grow inward and fuse into a single cavity that opens near the mouth on what is technically the dorsal area of the body.

External features
      A solitary tunicate has two major openings, or siphons, on the surface away from the area of attachment: a branchial aperture, through which water enters the body, and an atrial aperture, through which water, wastes, and gametes leave. Water circulation is produced by ciliary activity on the animal's pharynx. The animal is covered with a thick tunic, which consists of some cells, blood vessels, and a secretion of a variety of proteins and carbohydrates, including cellulose, which, although abundant in plants, is unusual in animals.

      Some solitary, sessile ascidians are stalked, and budding commonly occurs by growth at the base of the animal. In “social” colonial ascidians the zooids are relatively independent, whereas in “compound” colonial ascidians budding gives rise to a colony in which the zooids are embedded in a common tunic. Several zooids may share a single, common cloacal aperture through which water exits, but each zooid has its own branchial aperture through which water enters.

Internal features
Skeleton, tissues, and muscles
      The tunic functions to some extent as an external skeleton that supports and protects the body. Additional support is provided by body fluids and connective tissue. Firm proteinaceous rods also may support the branchial apparatus.

      Although musculature is poorly developed in tunicates, there are muscles that retract the body and constrict the atrial cavity, allowing it to eject water. In dolioloids and salps, these muscles have become modified so as to produce jet propulsion.

Nervous system and organs of sensation
      In the tadpole larvae and appendicularians, the dorsal nerve cord is well developed. At the anterior end there are usually sensory structures, which detect light and orient the animal to gravity. Similar sensory structures can be found in adult thaliaceans. Special organs of sense are otherwise poorly developed. When the larva metamorphoses into an adult, the original nervous system and sensory organs degenerate, leaving a single ganglion between the oral and atrial openings. Nerves grow to the various organs of the body from this ganglion.

      In ascidians and thaliaceans the beating action of pharyngeal cilia creates a water current. As the water is driven from the branchial sac into the atrial cavity, a sheet of mucus, secreted by the endostyle, traps a variety of very small organisms suspended in the water current, especially small plantlike protists (phytoplankton). The mucus is rolled into a cord and then conveyed to the intestine, where it is digested and absorbed. A stomach and glands may be present. The intestine ends as an anus in the atrium below the atrial aperture. Wastes are ejected through this aperture in a stream of water.

      Metabolic wastes, such as the breakdown products of protein, are excreted at various parts of the body, including the surfaces of the gills and the intestine, and sometimes by a discrete kidney. In many cases wastes are stored as solid deposits rather than being excreted from the body as they are produced.

      Gas exchange occurs across the gill and also across various other body surfaces, such as the lining of the atrium.

Water/vascular system
      Tunicates do not have the well-developed secondary body cavity (coelom) of other chordates, but traces of one perhaps are represented by cavities around the heart and by an extension of the gut called the epicardium around some of the internal organs. The body cavities are considered to be a part of the circulatory system. There are a heart and some large blood vessels but no tiny capillaries. The tunicate heart is unusual in that it periodically reverses the direction in which it pumps the blood, but the reasons for this behaviour are unknown. There are many different cell types in the blood.

Hormones
      A variety of possible endocrine organs help to coordinate feeding and reproduction. Various chemical substances are known to act as hormones in vertebrates; however, their exact role in the tunicates is uncertain.

Features of defense and aggression
      The tunic provides ascidians with some defense. They also may be protected by chemicals (such as sulfuric acid) that make them distasteful to predators. Appendicularians are small and therefore difficult to see. If attacked, they can escape from the house and form a new one. Thaliaceans are protected somewhat by transparency and can evade some predators by quickly ejecting a jet stream of water. They have well-developed light-producing organs, which may help to deter predators.

Evolution and paleontology
      Because they are soft-bodied animals, tunicates have left little fossil record apart from the hard mineral particles, called spicules, that are found in the tunics of some species. A single lineage within the class Ascidiacea, or perhaps a lineage of ascidian-like tunicates that branched off prior to the common ancestor of the Ascidiacea, probably gave rise to the other two classes. Embryonic thaliaceans show indications of having been derived from attached colonies. The pyrosomes, which resemble the colonies of some ascidians, evidently branched off first within the class Thaliacea and may not even be related to the dolioloids and salps. Appendicularians probably evolved from a more typical tunicate that reached sexual maturity before metamorphosis occurred. This development resulted in the loss of the adult stage (i.e., by paedomorphosis, retention of some juvenile features in the adult). Within the Ascidiacea, the common ancestor is generally thought to have been a solitary animal that did not reproduce by budding. The basis for this theory is that many ascidians do not bud, and the different patterns of budding that characterize distinct groups suggest independent origins. Evolution within the group has involved considerable elaboration of complex colonies, with the zooids themselves tending to become smaller and simpler in structure. There is a distinct trend toward parental care, especially in the colonial forms.

Classification

Annotated classification
Subphylum Tunicata (or Urochordata)
 Chordates with notochord restricted to the tail and, except in Appendicularia, only in tadpole larva; body covered with a tunic containing cellulose; atrium, except in Appendicularia, present and opening dorsally; heart present; coelom reduced; no clear traces of segmentation; about 2,600 species.
      Class Ascidiacea (sea squirts (sea squirt))
 Fixed as adults, solitary or colonial, oral and atrial apertures usually directed away from substrate; about 2,500 species.

      Subclass Enterogona
 Gonads unpaired, either within or behind intestinal loop; body may be divided into thorax and abdomen.

      Order Aplousobranchia
 Gills simple, unfolded and without longitudinal vessels or bars; digestive tract and genital organs in posterior part of body.

      Order Phlebobranchia
 Gills with longitudinal vessels and bars, without folds; gonads on one side, near digestive tract.

      Subclass Pleurogona
 Gonads and digestive tract by side of gill.

      Order Stolidobranchia
 Gill with longitudinal vessels, folded.

      Class Appendicularia (or Larvacea (larvacean))
 Adult small, pelagic, retaining larval notochord and tail; pharynx simple with two gill openings; no distinct atrium; about 70 species.

      Class Thaliacea
 Pelagic forms; atrial aperture directed toward the rear of each zooid; asexual buds form from a ventral stolon; about 70 species.

      Order Pyrosomida
 Zooids embedded in a tube open at one end.

      Order Doliolida
 Complex alternation of generations between a solitary, asexually and sexually reproducing gonozooid and colonial, asexually reproducing oozooids; gill with several to many stigmata.

      Order Salpida (salp)
 Complex alternation of generations between solitary, asexually reproducing oozooids and aggregated, sexually reproducing gonozooids. Pharynx leads to atrium by a single pair of slitlike openings; about 30 species.

Critical appraisal
      The above classification only approximates a natural, or genealogical, system. It is ambiguous with respect to the relationships of the three classes. Some authors put the class Appendicularia together with the class Thaliacea as Pelagotunicata, suggesting one possible relationship. Within the class Ascidiacea the system suggests that the original condition was that of the order Phelebobranchia and that the gill became simplified in a second group (order Aplousobranchia) and more complicated in a third (order Stolidobranchia). The Phlebobranchia, therefore, are not a single lineage but a grade, with some lineages close relatives of Aplousobranchia, others close relatives of Stolidobranchia, and others perhaps early branches. A change from simple to complex gills is also possible. Within the class Thaliacea the orders Doliolida and Salpida perhaps have a closer common ancestry with each other than with the order Pyrosomida, but this is not clear from the arrangement. Older systems were artificial rather than genealogical and often put all of the colonial forms together, even though it was known that colonial structure evolved more than once.

Michael T. Ghiselin

Additional Reading
E.J.W. Barrington, The Biology of Hemichordata and Protochordata (1965), is an account of the lower chordates and their evolution; N.J. Berrill, The Origin of Vertebrates (1955), argues the thesis that the urochordate larva represents the prototype from which cephalochordates and vertebrates are derived; A. Willey, Amphioxus and the Ancestry of the Vertebrates (1894), an early but good comprehensive account, presents the orthodox theory of chordate relationships; R.P.S. Jefferies, The Ancestry of the Vertebrates (1986), expounds an alternate theory of chordate origin; and Libbie H. Hyman, The Invertebrates, vol. 5, Smaller Coelomate Groups (1959), is a classic work treating the hemichordates in extensive detail. Later works include Charles K. Weichert and William Presch, Elements of Chordate Anatomy, 4th ed. (1975); and R. McNeill Alexander, The Chordates, 2nd ed. (1981); supplemented by Brian Bracegirdle and Patricia H. Miles, An Atlas of Chordate Structure (1978). N.J. Berrill, The Tunicata with an Account of the British Species (1950, reprinted 1968), a taxonomic survey with a useful section on tunicate biology; Pierre P. Grasse (ed.), Traité de zoologie: anatomie, systématique, biologie, vol. 11, Echinodermes, stomocordés, procordés (1966), an advanced zoological treatise devoted to protochordates, with good illustrations; W.A. Herdman, “Tunicata (Ascidians and Their Allies)” and “Cephalochordata,” in The Cambridge Natural History, vol. 7, pp. 35–138 (1904), an important general account; R.N. Millar, The Marine Fauna of New Zealand: Ascidiacea (1982), a morphological account of a single species of ascidian; Willard G. Van Name, “The North and South American Ascidians,” Bulletin of the American Museum of Natural History, vol. 84 (1945). For a later treatment, see “Invertebrate Chordates: Tunicates and Lancelets,” in Vicki Pearse et al., Living Invertebrates (1987).Michael T. Ghiselin

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Universalium. 2010.

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