chordate

/kawr"dayt/, Zool.

adj.

1. belonging or pertaining to the phylum Chordata, comprising the true vertebrates and those animals having a notochord, as the lancelets and tunicates.

n.

2. a chordate animal.

[1885-90; see CHORDATA]

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Any member of the phylum Chordata, which includes the most highly evolved animals, the vertebrates, as well as the marine invertebrate cephalochordates (see amphioxus) and tunicates.

All chordates, at some time in their life cycle, possess a dorsal supporting rod (notochord), gill slits, and a dorsal nerve cord. Unlike vertebrates, tunicates and cephalochordates lack any kind of brain or skeleton. Chordate bodies consist of a body wall encasing a gut, with a space between called the coelom. The body is usually long and bilaterally symmetrical, with the mouth and sense organs at the front end.

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▪ animal phylum

Introduction

      any member of the phylum Chordata, which includes the vertebrates, the most highly evolved animals, as well as two other subphyla—the tunicates and cephalochordates. Some classifications also include the phylum Hemichordata with the chordates.

      As the name implies, at some time in the life cycle a chordate possesses a stiff, dorsal supporting rod (the notochord). Also characteristic of the chordates are a tail that extends behind and above the anus, a hollow nerve cord above (or dorsal to) the gut, gill slits opening from the pharynx to the exterior, and an endostyle (a mucus-secreting structure) or its derivative between the gill slits. (A characteristic feature may be present only in the developing embryo and may disappear as the embryo matures into the adult form.) A somewhat similar body plan can be found in the closely related phylum Hemichordata.

General features

      Tunicates are small animals, typically one to five centimetres (0.4 to 2.0 inches) long, with a minimum length of about one millimetre (0.04 inch) and a maximum length slightly more than 20 centimetres; colonies may grow to 18 metres (59 feet) in length. Cephalochordates (cephalochordate) range from one to three centimetres. Vertebrates (vertebrate) range in size from tiny fish to the whales, which include the largest animals ever to have existed.

      Tunicates are marine animals, either benthic (bottom dwellers) or pelagic (inhabitants of open water), that often form colonies by asexual reproduction. They feed by taking water in through the mouth, using the gill slits as a kind of filter. The feeding apparatus in cephalochordates is similar. They have a well-developed musculature and can swim rapidly by undulating the body. Cephalochordates usually live partially buried in marine sand and gravel.

      Vertebrates retain traces of a feeding apparatus like that of tunicates and cephalochordates. The gill slits, however, ceased to function as feeding structures, and then later as respiratory devices, as the vertebrate structure underwent evolutionary changes. Except in some early branches of the vertebrate lineage (i.e., agnathans) a pair of gill arches has become modified so as to form jaws. The fishlike habitus that evidently began with cephalochordates became modified by the development of fins that were later transformed into limbs. With the invasion of the vertebrates into fresh water and then onto land, there was a shift in means of breathing—from gills to lungs. Other modifications, such as an egg that could develop on land, also emancipated the vertebrates from water. Elaboration of the locomotory apparatus and other developments allowed a diversification of structure and function that produced the amphibians, reptiles, birds, and mammals.

Natural history

Reproduction and life cycle

      The chordate life cycle begins with fertilization (the union of sperm and egg). In its primitive form, fertilization occurs externally, in the water. Asexual reproduction takes place in tunicates and in some vertebrates (females of some fish and lizards can reproduce without fertilization). Hermaphroditism (possessing both male and female reproductive organs) is found in tunicates and some fishes, but otherwise the sexes are separate. Larvae (very young forms that differ considerably from the juveniles and adults), when they do occur, differ in structure from the larvae of nonchordates. Internal fertilization, viviparity (giving birth to young that have undergone embryological development), and parental care are common in tunicates and vertebrates.

Ecology and habitats

      Chordates are common in all major habitats. Tunicate larvae either seek out a place where they can attach and metamorphose into an adult or develop into adults that float in the open water. Cephalochordates develop in the open water, but as adults they lie partially or entirely buried in sand and gravel. In either case, they are filter feeders with simple behaviour. Vertebrates are much more complex and, in keeping with their more active manner of obtaining food, highly varied in their ecology and habits.

Locomotion

      Chordates are capable of locomotion by means of muscular movements at some stage in life. In tunicate larvae, this is accomplished using a tail; in cephalochordates, by undulations of the body; and in vertebrates, by general body movements (as in eels and snakes) and by the action of fins and limbs, which in birds and some mammals are modified into wings.

Associations

      Chordates enter into a wide variety of symbiotic relationships and are especially noteworthy as hosts for parasites. Family groups and societal relationships, in both a broad and narrow sense, are particularly well developed in vertebrates, due primarily to their elaborate nervous systems. This phenomenon is seen in schools of fish, flocks of birds, and herds of mammals, as well as in the primate associations that suggest the beginnings of human society.

Form and function

General features

      Chordates have many distinctive features, suggesting that there has been extensive modification from simple beginnings. The early stages of chordate development show features shared with some invertebrate phyla, especially the mouth that forms separately from the anus, as it does in the phyla Hemichordata (hemichordate), Echinodermata, and Chaetognatha. Likewise, as in these phyla, the coelom, or secondary body cavity around the viscera, develops as outpouchings of the gut. A coelom also is present in some more distantly related phyla, including Annelida, Arthropoda, and Mollusca, but the main organs of the body are arranged differently in these phyla. In chordates the main nerve cord is single and lies above the alimentary tract, while in other phyla it is paired and lies below the gut. Cephalochordates and vertebrates are segmented, as are the annelids and their relatives; however, segmentation in the two groups probably evolved independently. The gill slits and some other features that are common among the hemichordates and the chordates originated before the chordates became a separate group. Hemichordates have no tail above the gut and no mucus-secreting endostyle between the gill slits.

External features

      An ancestral chordate, as suggested by the adult lancelet and the tadpole larva of tunicates, had a distinct front and hind end, an anterior mouth, a posterior tail above an anus, unpaired fins, and gill slits that opened directly to the exterior. A free-swimming tunicate larva metamorphoses into an attached, sessile adult with an atrium that surrounds the gills. The atrium of lancelets probably evolved independently.

Internal features

Skeleton and support

      The chordate notochord is a stiff rod with a turgid core and fibrous sheath. It keeps the animal from shortening when locomotory waves are produced through muscular contraction. The chordate body is supported by fluid in the body cavities. In tunicates, added support is provided by the tunic. Cartilaginous material supports the gills and other body parts of tunicates and cephalochordates. Immature vertebrate skeletons generally consist largely of cartilage, which becomes increasingly bony with age. The cartilaginous skeletons of sharks and some other vertebrates are thought to have evolved from more highly mineralized ones.

Tissues and muscles

      In both cephalochordates and vertebrates, muscles used in locomotion are well developed and organized segmentally. The tail musculature of tunicates is simpler and without clear indications of segmentation. There is at least a small amount of musculature throughout the body of all chordates. As jaws, limbs, and other body parts have evolved in vertebrates, so have the muscles that operate them.

Nervous system and sense organs

      The anterior end of the main nerve cord in chordates is enlarged to form at least the suggestion of a brain, but a brain is well developed only in vertebrates. Tunicate larvae have visual organs sensitive to light and sense organs responsive to the direction of gravity. Pigment spots and light receptors in the nerve cord of lancelets detect sudden changes in light intensity. The eyes and other sense organs of vertebrates are more elaborate and complex.

      The presence in cephalochordates and vertebrates of a nervous system with segmentally repeated nerves arising from the dorsal hollow nerve cord is suggestive of a common ancestry. The tunicate nervous system does not have the segmentally repeated nerves. The brains of all vertebrates are greatly enlarged and subdivided into functionally specialized regions.

Digestion and nutrition

      Both tunicates and cephalochordates are filter feeders of small particles of food suspended in the water. Beating cilia (hairlike cellular extensions) on the gill slits draw a current of water into the mouth and through the pharynx, where a sheet of mucus, secreted by the endostyle (a glandular organ lying below the two rows of gill slits), filters suspended food particles from the water. Cilia lining the pharynx move the food-rich sheet of mucus upward over the gill slits, and it is then rolled up and transported to the posterior part of the gut. The water current passes into the atrium and exits through the atrial opening.

      Something similar to this arrangement occurs in the vertebrates in the “ammocoetes” larva stage of the primitive jawless fish called the lamprey. The difference is that the food consists of somewhat larger particles that have been deposited on the bottom (detritus), and, instead of the feeding current being driven by cilia, the pharyngeal musculature pumps water and food particles across the gill slits. The earliest fishes probably fed on detritus, and a sucking action is retained by their extant representatives (lampreys and hagfishes). With the development of jaws, it became possible for the vertebrates to capture and seize larger food items.

      The lower digestive tract of the primitive chordate is a simple tube with a saclike stomach. There are only indications of the specialized areas and of glandlike structures, such as the liver and pancreas, that occur in vertebrates.

Excretion

      The excretion of wastes and the control of the chemical composition of the internal environment are largely effected by kidneys, although other parts of the body, including the gills, may play an important role. Tunicates and cephalochordates have a salt content essentially the same as seawater, but vertebrates, even marine species, have body fluids of low salt content, with the exception of hagfishes. A possible explanation is that the vertebrates evolved in fresh water, but it seems reasonable that hagfishes branched off while still marine and that the freshwater form evolved later.

Respiration

      A primitive chordate gill is present in tunicates and cephalochordates, where it serves in both respiration and feeding. The vertebrate gill may retain some role in feeding, although the current is now produced by the action of muscles, not cilia. The gills became reduced in number in various lineages, and they were strengthened by supporting elements, some of which evolved into jaws. Lungs, already present in fishes, became the main respiratory organs of terrestrial vertebrates.

Circulatory system

      The circulatory system in chordates has a characteristic pattern. In tunicates and vertebrates the blood is propelled by a distinct heart; in cephalochordates, by contraction of the blood vessels. Unoxygenated blood is driven forward via a vessel called the ventral aorta. It then passes through a series of branchial arteries in the gills, where gas exchange takes place, and the oxygenated blood flows to the body, much of it returning to its origin via a dorsal aorta. The blood of vertebrates passes through the tissues via tiny vessels called capillaries. In tunicates and cephalochordates, capillaries are absent and the blood passes through spaces in the tissues instead.

Hormones

      In vertebrates, endocrine glands (those of internal secretion) produce hormones that regulate many physiological activities. In tunicates and cephalochordates, organs have been identified that correspond in anatomical position to the pituitary gland of vertebrates, but which hormones, if any, they secrete is uncertain. In vertebrates, the thyroid gland produces thyroxine, an iodine-containing hormone that helps regulate metabolism. The thyroid is a modified endostyle, as can be illustrated by larval lampreys in which the thyroid still secretes mucus for use in feeding. The endostyles of lancelets take up iodine and form thyroxine, but the thyroxine formed may not function as a hormone in the lancelets themselves.

Features of defense and aggression

      Tunicates largely rely upon the passive defense afforded by their heavy tunic. Lancelets move rapidly through the substrate, and their well-developed locomotory apparatus evolved largely to provide a means of escaping predators. Vertebrates have ceased to feed on detritus brought to them by water currents. They have shifted to consuming larger foodstuffs and to actively locating, pursuing, and subduing what they eat.

Evolution and paleontology

      Chordates originated sometime earlier than 590 million years ago; that is, they predate the fossil record. Early representatives were soft-bodied, and they therefore left a poor fossil record. Fossils dubiously attributed to all three chordate subphyla have been found in Cambrian rocks (more than 505 million years old), and an extensive vertebrate fossil record begins around 400 million years ago.

      Embryological evidence places the phylum Chordata within the deuterostomes (Deuterostomia) (bilaterally symmetrical animals with undeterminate cleavage and whose mouth does not arise from the blastopore), which also includes the phyla Hemichordata, Echinodermata, and Chaetognatha. The closest relatives of the chordates are probably the hemichordates, since these animals possess gill slits and other features not found in other animal phyla. A slightly more remote relationship to the echinoderms is inferred on the basis of resemblances between the larvae in some groups of hemichordates and echinoderms. The derivation of chordates from certain fossil echinoderms has been argued on the basis of features such as what appear to be gill slits. Theories that derive them from other phyla (e.g., Annelida, Nemertea, Arthropoda) have been proposed, but such theories have few contemporary advocates.

      Whether the ancestral chordate was more like a tunicate or a cephalochordate has been extensively debated. The classical theory is that the ancestor was like a cephalochordate, and that one lineage became attached to hard surfaces and evolved into tunicates, whereas another remained unattached and evolved into vertebrates. An alternative theory is that the ancestor was like a tunicate and that the other two subphyla arose by modification of the tadpole larva. There is some preference for the classical theory because it provides the most satisfactory way of accounting for the similarities between chordates and hemichordates of the subphylum Enteropneusta. Within the chordates the tunicates probably branched off before the common ancestor of cephalochordates and vertebrates arose, for the latter resemble each other in some details of neuroanatomy and biochemistry.

Classification

Annotated classification

Phylum Chordata

 Deuterostomatous eucoelomates; gill clefts; endostyle or its derivative in pharynx; notochord; hollow dorsal nerve cord; tail posterior and dorsal to anus.

      Subphylum Tunicata (or Urochordata; tunicates (tunicate))

 Notochord, when present, restricted to tail; body covered with tunic, but sometimes only cuticle; atrium, absent in Appendicularia, dorsal and often paired in embryonic development; heart present; generally sessile (attached) as adults; see below Tunicates (tunicate).

      Class Ascidiacea (sea squirts)

 Sessile; benthic; solitary or colonial within a common tunic.

      Class Appendicularia (larvacea)

 Free-swimming; pelagic; resembles tadpole larvae of ascidians; 1 pair of gill slits; no distinct atrium.

      Class Thaliacea

 Pelagic; forms aggregations or colonies.

      Subphylum Cephalochordata (or Acrania; lancelets)

 Notochord extends entire body length, with tip anterior to nerve cord; atrium a single cavity with single, ventral opening; segments well developed; head poorly developed; no paired fins; no heart; see below Cephalochordates (cephalochordate).

      Subphylum Vertebrata (or Craniata; vertebrates)

 Notochord extends to the back of a well-developed head; no atrium; segments well developed; paired fins or limbs usually present; heart present; see below Vertebrates (vertebrate).

Critical appraisal

      This outline gives the major groups of chordates. Modern systematic biology attempts to arrange groups of organisms in a way that suggests the genealogical relationships (branching sequences) and therefore presents an epitome of evolutionary history. It also may attempt to show where there are important differences among the various groups. These goals often conflict. In a purely genealogical system, each group must correspond to a single lineage (clade) composed of the common ancestor and all of its descendants. A group that does not meet both of these requirements is called a grade and may be used as an informal group. Groups that do not contain the common ancestor, and therefore had two separate origins, are said to be polyphyletic. Such polyphyletic grades, which would put whales together with fish or birds together with bats, have generally been abandoned as soon as they were recognized. Another kind of grade, which does not include all the descendants of the common ancestor, is said to be paraphyletic and is retained in more conservative systems. Within the vertebrates the class Aves is a clade, but the class Reptilia is a grade, for the birds are modified dinosaurs. Some systems do not recognize Reptilia as a formal group. Likewise, birds, mammals, reptiles, and amphibians are all modified fish, and the old class of fishes (Pisces) is now rarely used. Vertebrata is a single clade, but “invertebrate” is a grade consisting of all animals except vertebrates. Therefore there is no formal group called Invertebrata.

      Many differences among systems are quite subjective. This is often the case when a group may be ranked either as a class or as a subphylum. The organizational limits of some groups are also largely a matter of opinion. Some authors have placed the phylum Hemichordata within the Chordata, expressing the close genealogical relationship. Others prefer to keep them as a separate phylum because hemichordates lack what are considered important chordate features.

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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