cephalochordate

/sef'euh loh kawr"dayt/, adj.

1. belonging or pertaining to the Cephalochordata.

n.

2. any chordate animal of the subphylum Cephalochordata, having fishlike characters but lacking a spinal column, comprising the lancelets.

[ < NL Cephalochordata; see CEPHALO-, CHORDATE]

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▪ chordate subphylum

Introduction

also called  acrania 

      any of more than two dozen species belonging to the subphylum Cephalochordata of the phylum Chordata. Small, fishlike marine invertebrates, they probably are the closest living relatives of the vertebrates. Cephalochordates and vertebrates have a hollow, dorsal nerve cord, pharyngeal gill slits, and a notochord. In most vertebrates, the embryonic notochord is eventually replaced by bony vertebrae or cartilaginous tissue; among cephalochordates, the notochord is retained into adulthood and is never replaced by vertebrae. There are about 20 species in two families, each with a single genus. Branchiostoma was formerly called Amphioxus, a name that is retained as an informal term. The other genus is Epigonichthys, also called Asymmetron. The genus Asymmetron is sometimes retained for some species. The cephalochordate fossil record extends back to about 525 million years ago during the early part of the Cambrian Period.

General features

Size and range of diversity of structure

      Adult lancelets reach a length of about six to seven centimetres (2.5 inches). There is little structural diversity within the group, the main difference between the two families being the restriction of the gonads to one side of the body in Epigonichthys.

Importance

      Although edible, lancelets are never sufficiently abundant to constitute a significant source of food to humans or an important part of the food chain in nature. Rather, their significance has to do with their place in evolution, as invertebrates transitional to vertebrates providing clues for the history of human lineage. This connection was first shown by the Russian zoologist Aleksandr Kowalevsky in 1867 in embryological evidence that was influential in establishing that evolution has in fact occurred. More recently, the relationship has been well-supported by gene sequence comparisons. Lancelets have a structure that illustrates the characteristic features of chordates in simple form.

Natural history

Reproduction and life cycle

      Lancelet sexes are separate, and asexual reproduction does not occur. Eggs and sperm are shed directly into the water, where fertilization occurs. The early stages of development strikingly resemble those of both tunicates and vertebrates. A larva is produced that is similar in structure to the adult but is peculiarly asymmetrical (the gill slits on one side develop first), smaller, and simpler, with fewer gill slits and no atrium. The larvae spend much of their time feeding in the open water but can be found on the bottom. After growing and developing, they metamorphose into the adult form and complete their life history in the substrate.

Ecology and habitats

      Lancelets are distributed throughout the world along tropical and temperate coasts. They inhabit soft bottoms ranging from sand to coarse shelly sand or gravel in shallow coastal water. Lancelets lie buried beneath this substrate, often with their mouths protruding above the surface, allowing them to take in water laden with food. In China, lancelets are sometimes eaten and even support a small fishing industry.

Food, feeding, and movement

      Lancelets can swim both forward and backward and can move rapidly through the gravel in which they live. Their behaviour is simple, largely being a matter of locating the proper habitat and escaping from predators. Larvae filter small organisms out of the water; at the time when they metamorphose into the adult, they also feed upon coarser materials deposited on the bottom. The adults filter small organisms from the overlying water by drawing a current into the mouth. The tentacle-like cirri around the mouth form a grid that keeps out sand and other large particles.

Form and function

General features

      The lancelets are also called cephalochordates (Greek: kephale, “head”) because the notochord extends from near the tip of the tail to well into the anterior of the body. Because they do not have the braincase, or cranium, of a vertebrate, lancelets are often called acraniates. The pharynx, with its many gill slits, is surrounded by the atrium, a large cavity with a single exit (the atriopore) on the lower surface of the body. The atrium protects the gills. Tunicates also have an atrium, but its evolution is probably independent of that of the cephalochordrate atrium.

      The bodies of lancelets, like those of fishes and other vertebrates, are largely made up of serially repeated units (segments) that include blocks of muscles called metameres. This segmentation also extends to the nerves that supply the myotomes and to some body cavities, excretory structures, and other parts. Segmentation is thought to provide more effective body coordination during locomotion. The segments of vertebrates and cephalochordates are so similar that they were almost certainly present in the common ancestor of the two groups. Tunicates and hemichordates have no clear indications of ever having possessed segments. Segments occur in other animals, including annelid worms and arthropods, but these segments have a different composition and probably a separate evolutionary origin.

      A distinct “secondary” body cavity (coelom), like that which contains the internal organs in vertebrates and many other animals, is well developed and forms a system of cavities and spaces. Like the coelom of hemichordates, echinoderms, and a few other animals, it develops as outpouchings in the gut of the embryo.

External features

      Lancelets are streamlined animals. A dorsal fin extends along the upper surface of the body and continues as a caudal fin around a tail and as a ventral fin to an atrium on the lower surface. Paired fins are absent, but metapleural folds along the sides of the body suggest precursors of paired fins. The tip of the body projects slightly above and in front of the mouth, which is surrounded by a funnellike oral hood that bears the cirri. The anus opens well behind the atriopore, on the left side of the ventral fin. The general body surface is covered by a smooth cuticle layer.

Internal features

Skeleton, tissues, and muscles

      The notochord extends virtually the entire length of the body and provides much of its support. It has a firm sheath and a core made up of a single series of cells that contains muscle fibres. These fibres probably maintain the stiffness of the notochord, the main role of which is keeping the body from shortening when the animal swims. The gills, fins, and cirri also contain stiff, supportive rods.

      The main body musculature occurs in horizontal chevron-shaped blocks of muscle (myotomes) like those of fishes. This arrangement allows the muscles to pull more effectively in producing a side-to-side movement of the body in swimming. The remaining muscles are quite small and associated largely with feeding and the movement of internal organs.

Nervous system and organs of sensation

      The cephalochordate nervous system is simple. The main nerve cord, which is single and hollow as in all chordates, has a slight swelling at the front that barely qualifies as a brain. Nerves from the main nerve cord occur in groups that roughly compare to those of vertebrates in arrangement and in the regions supplied. There are small eyelike organs in the nerve cord that can detect the direction of light and changes in its intensity. Various areas of the body surface, including some near the mouth, detect chemicals in the water and thereby aid in feeding.

Digestion and excretion

      Lancelets are suspension feeders that extract small particles suspended in the water. The mouth is covered by an oral hood, the edges of which form the buccal cirri. The cephalochordate commonly is buried in the substrate and positions its mouth above the surface of the sand. During feeding, the cirri form a kind of grid that keeps out large particles. Water is drawn into the mouth by the beating action of cilia on the gills. The pharynx is a large section of the gut just behind the mouth, extending about two-thirds the length of the body, with many narrow gill slits. The water current enters the pharynx, passes through the gill basket to the atrium, and leaves the body through the atriopore. On the floor of the pharynx, between the left and right series of gill slits, an endostyle secretes a sheet of mucus that moves upward along the gills and traps food particles suspended in the water current. The mucus is rolled up and transported to the intestine, where food is digested and absorbed. There is no distinct stomach. The intestine is straight, except for a blind outpouching called the caecum (cecum), which has, on the basis of position, been compared to the liver and pancreas of vertebrates. It extends forward along the right side of the pharynx.

      Lancelets have unique excretory structures called solenocytes, which occur only in some distantly related animals, such as annelids.

Respiration

      The gill is largely a feeding organ, but it also serves for the exchange of gases in respiration. After the water has passed through the gill slits, it reaches the atrium and exits through the atriopore. Excretory products and eggs and sperm also exit the body through this opening.

Circulatory system

      The general pattern of blood circulation through vessels and tissues in cephalochordates is strikingly like that of vertebrates, although simpler. The most notable difference is that cephalochordates lack a heart. Blood is forced through the closed system by contractile blood vessels (especially one called the ventral aorta) and by blood vessels of the gills. Blood passes forward from the rear of the body to the ventral aorta, which is located beneath the endostyle, and then branches upward through vessels in the gills. Most of the blood then passes toward the rear of the animal, some of it moving through capillaries in the intestine and taking up food. From the posterior end of the body, blood passes forward and then makes a detour through capillaries in the caecum, much as it does through the liver of lower vertebrates, back to the ventral aorta. There are no corpuscles in the blood.

Hormones

      The endostyle takes up iodine and forms thyroxine, an important hormone produced by the vertebrate thyroid gland. This homology is interpreted as a step in the evolution of the thyroid from the endostyle. It is not certain what role thyroxine plays in the physiology of the lancelets themselves, however.

Evolution and paleontology

      Soft-bodied animals such as lancelets rarely have a good fossil record. A few fossils have been interpreted as cephalochordates, but few of these determinations are well founded. A good possibility is Pikaia, a fossil discovered in the Burgess Shale (Middle Cambrian, about 530 million years old). Pikaia has myotomes and what looks like a notochord, indicating that it is a chordate, but only its shape suggests that it is a lancelet rather than a fish.

      The cephalochordates make plausible models for the common ancestor of the chordates and for precursors to vertebrates. They evidently represent a collateral branch on the vertebrate lineage that has been somewhat modified since common ancestry, however, and should not be thought of as a human ancestor. Several features unique to cephalochordates and vertebrates suggest that they are “sister groups” more closely related to each other than either is to other chordates. These features include the segmented musculature and its innervation, the pattern of circulation, and several biochemical features. The atrium is thought to have evolved independently in cephalochordates and tunicates; hence there is little evidence for the two forming a single lineage.

      Whether the ancestral chordate was more like a cephalochordate or a tunicate is debatable, because features absent in tunicates could mean that they never have been present or could mean that they have been lost. Some authors have argued that the simplicity of cephalochordates is due to degeneration, but there are no clear indications that this is true.

Classification

Phylum Chordata

 

      Subphylum Cephalochordata

 Notochord extends anterior to dorsal nerve cord; atrium ventral; segments and coelom well developed; no heart; no paired fins; more than 24 species.

      Family Branchiostomatidae

 Double row of gonads; Branchiostoma.

      Family Epigonichthyidae

 Gonads on right side of body only; Epigonichthys.

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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