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AMPHIOXUS, or LANCELET

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Originally appearing in Volume V01, Page 891 of the 1911 Encyclopedia Britannica.
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AMPHIOXUS, or LANCELET, the name of small, fish-like, marine creatures, forming the class Cephalochorda, of the phylum Vertebrata. Lancelets are found in brackish or salt water, generally near the coast, and have been referred to several genera and many species. They were first discovered by P. S. Pallas in 1778, who took them to be slugs and described them under the name Limax lanceolatus. The true position in the animal kingdom was first recognized in 1834 by O. G. Costa, who named the genus Branchiostoma, and it has since been dealt with by many writers. The theoretical interest of Amphioxus depends upon a variety of circumstances. In its manner of development from the egg, and in the constitution of its digestive, vascular, respiratory (branchial), excretory, skeletal, nervous and muscular systems it exhibits what appears to be a primordial condition of vertebrate organization, a condition which is, in fact, partly recapitulated in the course of the embryonic stages of craniate vertebrates. In comparative morphology it provides many illustrations of important biological principles (such, for example, as substitution and change of function of organs), and throws new light upon, or at least points the way to new ideas of, the primitive relations of different organic systems in respect of their function and topography. One of the most puzzling features in its structure, and, at the same time, one of the greatest obstacles to the view that it is essentially primitive and not merely a degenerate creature, is the entire absence of the paired organs of special sense, olfactory, optic and auditory, which are so characteristic of the higher vertebrates. Although it is true that there is a certain amount of gradation in the degree of development to which these organs have attained in the various orders, yet it is hardly sufficient to enable the imagination to bridge over the gap which separates Amphioxus from the lowest fishes in regard to this feature of organization. Classification.—On account of the absence of anything in the nature of a skull, Amphioxus has been regarded as the type of a division, Acrania, in contrast with the Craniata which comprise all the higher Chordata. The ordinal name for the genera and species of Amphioxus is Cephalochorda, the term referring to the extension of the primary backbone or notochord to the anterior extremity of the body; the family name is Branchiostomidae. The amount of generic divergence exhibited by the members of this family is not great in the mass, but is of singular interest in detail. There are two principal genera—r. Branchiostoma Costa, having paired sexual organs (gonadic pouches); 2. Heteropleuron Kirkaldy, with unilateral gonads. Of these, the former includes two subgenera, Amphioxus (s. str.) Yarrell and Dolichorhynchus Willey. The species belonging to the genus Heteropleuron are divided among the three subgenera Paramphioxus Haeckel, Epigonichthys Peters, and Asymmetron Andrews. The generic characters are based upon definite modifications of form which affect the entire facies of the animals, while the specific diagnoses depend upon minor characters, such as the number of myotomes or muscle-segments. Habits and Distribution.—With regard to its habits, all that need be said here is that while Amphioxus is an expert swimmer when occasion requires, yet it spends most of its time burrowing in the sand, in which, when at rest, it lies buried with head protruding and mouth wide agape. Its food consists of microscopic organisms and organic particles; these are drawn into the mouth tm i rgo 02 20 t l an together with currents of water induced by the action of the vibratile cilia which are abundant along special tracts on the sides and roof of the vestibule of the mouth and in the walls of the perforated pharynx ("ciliary ingestion"). Amphioxus favours a littoral habitat, and rarely if ever descends below the 50-fathom line. Species occur in all seas of the temperate, tropical and subtropical zones. The European species. A. lanceolatus, is found in the Black and Mediterranean Seas, and on the coasts of France, Great Britain and Scandinavia, while a closely allied species or subspecies, A. caribaeus, frequents the Caribbean region from Chesapeake to La Plata. A. californiensis occurs on the coast of California, and A. belcheri extends its area of distribution from Queensland through Singapore to Japan. A recently described species, Dolichorhynchus indicus, characterized by the great length of the praeoral lobe or snout, has been dredged in the Indian Ocean, Paramphioxus bassanus occurs on the coast of Australia from Port Phillip to Port Jackson; P. cingalensis at Ceylon. Epigonichlhys cultellus (fig. I) inhabits Torres Strait, and has also been body (these have no constant numerical relation to the myomeres); M, notochord as seen through the transparent myotomes, the thin double-lined spaces being the connective-tissue septa and the broader spaces the muscular tissue of the myotomes; N, position of brown funnel of left side (atrio-coelomic canal); 0, nerve tube resting on notochord. (2) Dissection of Amphioxus. By a horizontal incision on each side of the body a large ventral area has been separated and turned over, as it were on a hinge, to the animal's left side. The perforated pharyngeal region has then been detached from the adherent epipleura or opercular folds (wall of atrial or branchial chamber) by cutting the fluted pharyngo-pleural membrane d, and separated by a vertical' cut from the intestinal region. a, Edge of groove formed by adhesion of median dorsal surface of alimentary canal to sheath of notochord; b, median dorsal surface of alimentary canal; c, left dorsal aorta; cc, single dorsal aorta, formed by union of the two anterior vessels; cc', same vessel resting on intestine; d, cut edge of pharyngo-pleural folds of atrial tunic, really the original outer body-wall before the downgrowth of epipleura; d', atrial tunic (original body-wall) at non-perforate region, cut and turned back so as to expose peri-enteric coelom and intestine r; e', upstanding folds of body-wall (pharyngo-pleural folds) on alternate bars of perforate region of body; f, atrio-coelomic canals or brown funnels (collar-pores of Balanoglossus); g, cavity of a gonad-sac; m, cut musculature of body-wall; n, anus; o, post-atrioporal extension of atrial chamber in form of a tubular caecum; p, atriopore; q, hepatic caecum; r, intestine; s, coelom; t, area of adhesion between alimentary canal and sheath of notochord; v, atrial chamber or branchial cavity; w, post-atrioporal portion of intestine; x, canals of metapleura exposed by cutting; E, probe passing through atriopore into atrial or branchial chamber; FF , probe passing from coelom, where it expands behind the atriopore, into narrower perienteric coelom of praeatrioporal region. (3) Portion of (2) enlarged to show atrio-coelomic canals ("brown funnels " of Lankester). Lettering as in (2). (¢) Section taken transversely through praeoral region near termination of nerve tube. a, Olfactory ciliated pit on animal's left side, its wall confluent with substance of nerve tube; b, pigment spot (rudimentary eye) on anterior termination of nerve tube; c, first pair of nerves in section; d, fin ray; e, myotome; f, notochord; g, space round myotome (?artifact or coelom) ; h, subchordal canal (? blood-vessel); i, a symmetrical epipleura of praeoral hood. found at Ternate. Asymmetron lucayanum is the Bahaman representative of the family, with a subspecies, A. caudatum, in the South Pacific from New Guinea to the Loyalty Islands. The Peruvian species, Branchiostoma elongatum, with nearly eighty myotomes, cannot at present be assigned to its proper subgenus. External Form.—The following description, unless otherwise stated, refers to A. lanceolatus. Amphioxus is a small fish-like creature attaining a maximum length of about 3 in., semi-transparent in appearance, showing iridescent play of colour. The body is narrow, laterally compressed and pointed at both ends. The main musculature can be seen through the thin skin to be divided into about sixty pairs of muscle-segments (myotomes) by means of comma-shaped dissepiments, the myocommas, which stretch between the skin and the central skeletal axis of the body. These myotomes enable it to swim rapidly with characteristic serpentine undulations of the body, the movements being effected by the alternate contraction and relaxation of the longitudinal muscles on both sides. Apparently correlated with this peculiar locomotion is the anatomical fact of the alteration of the myotomes on the two sides. Symmetrical at their first appearance in the embryo, the somites (from which the myotomes are derived) early undergo a certain distortion, the effect of which is to carry the somites of the left side forwards through the length of one half-segment. For example, the twenty-seventh myotome of the left side is placed opposite to the twenty-sixth myocomma of the right side. The back of the body is occupied by a'crest, called the dorsal fin, consisting of a hollow ridge, the cavity of which is divided into about 250 compartments or fin chambers, into each of which, with the exception of those near the anterior and posterior end of the body, projects a stout pillar composed of characteristic laminar tissue, the fin ray. The dorsal crest is continued round both extremities, becoming expanded to form the rostral fin in front and the caudal fin behind. Even in external view, careful inspection will show that the body is divisible into four regions, namely, cephalic, atrial, abdominal and caudal. The cephalic region includes the rostrum or praeoral RIGHT lobe and the mouth. As already stated, the notochord extends beyond the mouth to the tip of the rostrum. The mouth consists of two portions, an outer vestibule and an inner apertura oris; the latter is surrounded by a sphincter muscle, which forms the so-called velum. The vestibule of the mouth is the space bounded by the oral hood; this arises by secondary downgrowth of lid-like folds over the true oral aperture, and is provided with a fringe of tentacular cirri, each of which is supported by a solid skeletal axis. The oral hood with its cirri has a special nerve supply and musculature by which the cirri can be either spread out, or bent inwards so that those of one side may interdigitate with those of the other, thus completely closing the en-trance to the mouth. The velum is also provided with a circlet of twelve tantacles (in some species sixteen) which hang backwards into the pharynx; these are the velar tentacles. The atrial region extends from the mouth over about two-thirds of the length of the body, terminating at a large median ventral aperture, the atriopore; this is the excurrent orifice for the respiratory current of water and also serves for the evacuation of the generative products. This region is really the branchiogenital region, al-though the fact is not apparent in externaI view. The ventral side of the body in the atrial region is broad and convex, so that the body presents the appearance of a spherical triangle in transverse section, the apex being formed by the dorsal fin and the angles bordered by two hollow folds, the metapleural folds, each of which contains a continuous longitudinal lymph-space, the metapleural canal. In the genus Branchiostoma the meta-pleural folds terminate symmetrically shortly behind the atriopore, but in Heteropleuron the right metapleur passes uninterruptedly into the median crest of the ventral fin (fig. I). In this connexion it may also be mentioned that in all cases the right half of the oral hood is directly continuous with the rostral fin (fig. 2). The abdominal region comprises a short stretch of body between atriopore and anus, the termination of the alimentary canal. It is characterized by the presence of a special development of the lophioderm or median fin-system, namely, the ventral fin, which is composed of two portions, a lower keel-like portion, which underlies an upper chambered portion, each chamber containing typically a pair of gelatinous fin rays. Finally, the caudal region comprises the post-anal division of the trunk. The keel of the ventral fin is continued past the anus into the expanded caudal fin, and so it happens that the anal opening is displaced from the middle line to the left side of the fin. In Asymmetron the caudal region is remarkable for the curious elongation of the notochord, which is produced far beyond the last of the myotomes. Alimentary, Respiratory and Excretory Systems.—Although the function of the two latter systems of organs is the purification of the blood, they are not usually considered together, and it is therefore the more remarkable that their close association in Amphioxus renders it necessary to treat them in common. The alimentary canal is a perfectly straight tube lined throughout by ciliated epithelium. As food particles pass in through the mouth they become enveloped in a slimy substance (secreted by the endostyle) and conveyed down the gut by the action of the vibratile cilia as a continuous food-rope, the peristaltic movements of the gut-wall being very feeble. The first part of the alimentary canal consists of the pharynx or branchial sac, the side walls of which are perforated by upwards of sixty pairs of elongatedslits, the gill-clefts. Each primary gill-cleft becomes divided into two by a tongue-bar which grows down secondarily from the upper wall of the cleft and fuses with the ventral wall. New clefts continue to form at the posterior end of the pharynx during the adult life of the animal. The gill-clefts open directly from the cavity of the pharynx into that of the atrium, and so give egress to the respiratory current which enters the mouth with the food (fig. 4). The atrium or atrial chamber is a peripharyngeal cavity of secondary origin effecting the enclosure of the gill-clefts, which in the larva opened directly to the exterior. The atrium is thus analogous to the opercular cavity of fishes and tadpoles, and, as stated above, remains in communication with the exterior by means of the atriopore. The primary and secondary bars which separate and divide the successive gill-clefts from one another are traversed by blood-vessels which run from a simple tubular con-tractile ventral branchial vessel along the bars into a dorsal aorta. The ventral branchial vessel lies below the hypobranchial groove or endostyle, and is the representative of a heart. As water for respiration streams through the clefts, gaseous interchange takes place between the circulating colourless blood and the percolating water. The pharynx projects freely into the atrium; it is surrounded at the sides and below by the continuous atrial cavity, but dorsally it is held in position in two ways. First, its dorsal wall (which is grooved to form the hyperpharyngeal groove) is closely adherent to the sheath of the notochord; and secondly, the pharynx is attached through the intermediation of the primary bars. These are suspended to the muscular body-wall by a double membrane, called the ligamentum denticulatum, which forms at once the roof of the atrial chamber and the floor of a persistent portion of the original body-cavity or coelom (the dorsal coelomic canal on each side of the pharynx). The ligamentum denticulatum is thus lined on one side by the epiblastic atrial epithelium, and on the other by mesoblastic coelomic epithelium. Now this ligament is inserted into the primary bars some distance below the upper limits of the gill-clefts, and it therefore follows that, corresponding with each tongue-bar, the atrial cavity is produced upward beyond the insertion of the ligament into a series of bags or pockets, which may be called the atrial pouches. At the top of each of these pouches there is a minute orifice, the aperture of a small tubule lying above each pouch in the dorsal coelom. These tubules are the excretory tubules or nephridia. They communicate with the coelom by several openings or nephrostomes, and with the atrium by a single opening in each case, the nephridiopore. It is important to emphasize the fact that in Amphioxus the excretory tubules are co-extensive with the gill-clefts. The perforated pharynx terminates some distance in front of the atriopore. At the level of its posterior end a pair of funnel-shaped pouches of the atrium are produced forwards into the dorsal coelom. These are the atrial coelomic funnels or brown funnels, so called on account of the characteristic pigmentation of their walls. There are reasons for supposing that these funnels are vestiges of aIi ancient excretory system, which has given way by substitution to the excretory tubules described above. In the same region of the body, namely, close behind the pharynx, a large diverticulum is given off from the ventral side of the gut. This is the hepatic caecum (fig.z,z, q, fig. 4, 1), which is quite median at its first origin, but, as it grows in length, comes to lie against the right wall of the pharynx. Although within the atrial cavity, it is separated from the latter by a narrow coelomic space, bounded towards the atrium by coelomic and atrial epithelium. No food passes into the hepatic caecum, which has been definitely shown on embryological and physiological grounds to be the simplest persistent form of the vertebrate liver. Nervous System.—As has already been indicated, a solid sub-cylindrical elastic rod, the notochord, surrounded by a sheath of laminar connective tissue, the cordal sheath, lies above the alimentary canal in contact with its dorsal wall, and extends beyond it both in front and behind to the obtusely pointed extremities of the body. This notochord represents the persistent primordial skeletal axis which, in the higher Craniata (though not so in the lower), gives way by substitution to' the segmented vertebral column. Immediately above the notochord there lies another elongated slit when relaxed (as in the lamprey) ; p, perforated pharynx; e, endostyle; g, gonads; 1, liver; at, level of atriopore; i, intestine; an, anus. In this species the atrium is produced as an asymmetrical blind pouch behind the atriopore as far as the anus. subcylindrical cord, also surrounded by a sheath of connective tissue. This cord is neither elastic nor solid, but consists of nerve tissue, fibres and ganglion cells, surrounding a small central canal. For the sake of uniformity in nomenclature this nerve-cord may he called the neurochord. It is the central nervous system, and contains within itself the elements of the brain and spinal marrow of higher forms. The neurochord tapers towards its posterior end, where it is coextensive with the notochord, but ends abruptly in front, some distance behind the tip of the snout. The neurochord attains its greatest thickness not at its anterior end but some way behind this region; but the central canal dilates at the anterior extremity to form a thin-walled cerebral vesicle, in the front wall of which there is an aggregation of dark pigment cells constituting an eyespot, visible through the transparent skin (fig. I). There are two pairs of specialized cerebral nerves innervating the praeoral lobe, and provided with peripheral ganglia placed near the termination of the smaller branches. Corresponding with each pair of myotomes, and subject to the same alternation, two pairs of spinal nerves arise from the neurochord, namely, a right and left pair of compact dorsal sensory roots without ganglionic enlargement, and a right and left pair of ventral motor roots composed of loose fibres issuing separately from the neurochord and passing directly to their termination on the muscle-plates of the myotomes. The first dorsal spinal nerve coincides in position with the myocomma which separates the first myotome from the second on each side, and thereafter the successive dorsal roots pass through the substance of the myocommata on their way to the skin; they are therefore septal or intersegmental in position. The ventral roots, on the contrary, are myal or seg- mental in position. In addition to the cerebral eyespot there are large numbers of minute black pigmented bodies beside and below the central canal of theneuro- chord, commencing from the level of the third myotome. It has been determined that these bodies are of the nature of eyes (Becheraugen, R. Hesse), each consisting of two cells, a cup-shaped pigment cell and a triangular retinal cell. These may be called the spinal eyes, and it is said that they are disposed in such a way as to receive illumination preferentially from the right side, although this fact has no relation with the side upon which Amphi- oxus may lie upon the sand. When kept in captivity the animal often lies upon one side on the surface of the sand, but on either side indiffer- ently. Over the cerebral eye there is a small orifice placed to the left of the base of the cephalic fin, leading into a pit which extends from the surface of the body to the surface of the cerebral vesicle; this is known as A. von Killiker's olfactory pit. Reproductive System.—The sexes are separate, and the male or female gonads, which are exactly similar in outward appearance, occur as a series of gonadic pouches projecting into the atrial cavity at the base of the myotomes (figs. 2, 3, 4). At the breeding season the walls of the pouches burst and the sexual elements pass into the atrium, whence they are discharged through the atriopore into the water, where fertilization takes place. Development.—The development of Amphioxus possesses many features of interest, and cannot fail to retain its importance as an introduction to the study ofembryology. The four principal phases in the development are: (1) Blastula, (2) Gastrula, (3) Flagellate Embryo, (4) Larva. The segmentation or cleavage of the ovum which follows upon fertilization terminates in the achievement of the blastula form, a minute sphere of cells surrounding a central cavity. Then follows the phenomenon of gastrulation, by which one-half of the blastula is invaginated into the other, so as to obliterate the segmentation cavity. The embryo now consists of two layers of cells, epiblast and hypoblast, surrounding a cavity, the archenteron, which opens to the exterior by the orifice of invagination or blastopore. One important fact should IliNlll illfln'"NNNMICryIIW,ApII111nIXm null wIII1R nNNN aluenae—a,.npl lit m 77k,gt lips figure.—anp, Anterior neural pore; bc, rudiment of buccal skeleton; c, cilia; cb, ciliated band; cc, ciliated groove; cm, cilia at margin of mouth ; gl, external opening of club-shaped gland ; Hn, Hatschek's nephridium; lm, left metapleur; n, notochord; pp, praeoral pit; ps, primary gill-slits, 1, 5, and 13; rm, right metapleur showing through. Lettering of lower figure.—a, Atrium ; at, alimentary canal; bv, blood-vessel; cv, cerebral vesicle; df, dorsal section of myocoel (= fin spaces) ; e, " eyespot "; end, endostyle; gl, club-shaped gland; lm, edge of left metapleur; m, lower edge of mouth; n, notochord; nt, pigmented nerve tube; ps, primary gill-slits, 1, 9, and 14; rc, renal cells on atrial floor; rm, edge of right metapleur; so, sense organ opening into praeoral pit ; ss, thickenings, the rudiments of the row of secondary gill-slits. be noted with regard to the gastrula, in which it seems to differ from the gastrulae of invertebrata. After invagination is completed, the embryo begins to elongate, the blastopore becomes narrower, and the dorsal wall of the gastrula loses its convexity, and becomes flattened to form the dorsal plate, the outer layer of which is the primordium of the neurochord and the inner layer the primordium of the notochord. While still within the egg-membrane the epiblastic cells become flagellated, and the gastrula rotates within the membrane. About the eighth hour after commencement of development the membrane ruptures and the oval embryo escapes, swimming by means of its flagella at the surface of the sea for another twenty-four hours, during which the principal organs are laid down, although the mouth does not open until the close of this period. The primordium of the neurochord (neural or medullary plate) referred to above becomes closed in from the surface by the overgrowth of surrounding epiblast, and its edges also bend up, meet, and finally fuse to form a tube, the medullary or neural tube. An important fact to note is that the blastopore is included in this overgrowth of epiblast, so that the neural tube remains for some time in open communication with the archenteron by means of a posterior neurenteric canal. It is still longer before the neural tube completes its closure in front, exhibiting a small orifice at the surface, the anterior neuropore. It is thus possible that the neurenteric canal is due to the con-junction of a posterior neuropore with the blastopore, i.e. it is a complex and not a simple structure. Paired archenteric pouches m llmu nN ,u~u motmiont!T!9nu ml uml ummutm llINIIIINIIIIMHOMHR tl I m NWM meanwhile appear at the sides of the axial notochordal tract, the mesoblastic somites. The first of these differs in several respects from those which succeed, and has been called the collar cavity (MacBride). In front of the latter there remains a portion of the archenteron, which becomes constricted off as the head cavity. This becomes divided into two, the right half forming the cavity of the rostrum, while the left acquires an opening to the exterior, and forms the praeoral pit of the larva, which subsequently gives rise to special ciliated tracts in the vestibule of the mouth mentioned above. The larval period commences at about the thirty-sixth hour with the perforation of the mouth, first gill-cleft and anus. The larva is curiously asymmetrical, as many as fourteen gill-clefts appearing in an unpaired series on the right side, while the mouth is a large orifice on the left side, the anus being median. The adult form is achieved by metamorphosis, which cannot be further described here. One point must not be omitted, namely, the homogeny of the endostyle of Amphioxus and the thyroid gland of Craniata. and the Ancestry of the Vertebrates (1894) ; Remarks on some recent Work on the Protochorda," Quart. J. Micr. Sci. xxiii. (1899), p. 223; " Dolichorhynchus indicus," ib. (1901); W. B. Benham, Heteropleuron of New Zealand," ib. (1901) E Burchardt, " Finer Anatomy of Amphioxus," with bibliography, Jena Zeitschr. xxxiv. (1900), P. 719. (A. W.`)
End of Article: AMPHIOXUS, or LANCELET
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