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AMPHIOXUS, or LANCELET
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See also:AMPHIOXUS, or LANCELET , the name of small, See also:fish-like, marine creatures, forming the class Cephalochorda, of the phylum See also:Vertebrata. Lancelets are found in brackish or See also:salt See also:water, generally near the See also:coast, and have been referred to several genera and many See also:species. They were first discovered by P. S. See also:Pallas in 1778, who took them to be slugs and described them under the name Limax lanceolatus. The true position in the See also:animal See also:kingdom was first recognized in 1834 by O. G. See also:Costa, who named the genus Branchiostoma, and it has since been dealt with by many writers. The theoretical See also:interest of Amphioxus depends upon a variety of circumstances. In its manner of development from the See also:egg, and in the constitution of its See also:digestive, vascular, See also:respiratory (branchial), excretory, skeletal, See also:nervous and See also:muscular systems it exhibits what appears to be a primordial See also:condition of vertebrate organization, a condition which is, in fact, partly recapitulated in the course of the embryonic stages of craniate vertebrates. In See also:comparative See also:morphology it provides many illustrations of important biological principles (such, for example, as substitution and See also:change of See also:function of See also:organs), and throws new See also:light upon, or at least points the way to new ideas of, the See also:primitive relations of different organic systems in respect of their function and See also:topography. One of the most puzzling features in its structure, and, at the same See also:time, one of the greatest obstacles to the view that it is essentially primitive and not merely a degenerate creature, is the entire See also:absence of the paired organs of See also: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 See also:imagination to See also:bridge over the See also:gap which separates Amphioxus from the lowest fishes in regard to this feature of organization. See also:Classification.—On See also:account of the absence of anything in the nature of a See also:skull, Amphioxus has been regarded as the type of a See also:division, See also: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 See also:term referring to the See also:extension of the See also:primary backbone or notochord to the anterior extremity of the See also:body; the See also:family name is Branchiostomidae. The amount of generic divergence exhibited by the members of this family is not See also:great in the See also:mass, but is of singular interest in detail. There are two See also: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.) See also:Yarrell and Dolichorhynchus Willey. The species belonging to the genus Heteropleuron are divided among the three subgenera Paramphioxus See also:Haeckel, Epigonichthys See also:Peters, and Asymmetron See also:Andrews. The generic characters are based upon definite modifications of See also:form which affect the entire facies of the animals, while the specific diagnoses depend upon See also:minor characters, such as the number of myotomes or muscle-segments. Habits and See also:Distribution.—With regard to its habits, all that need be said here is that while Amphioxus is an See also:expert swimmer when occasion requires, yet it spends most of its time burrowing in the See also:sand, in which, when at See also:rest, it lies buried with See also:head protruding and mouth wide See also:agape. Its See also:food consists of microscopic organisms and organic particles; these are See also:drawn into the mouth tm i rgo 02 20 t l an together with currents of water induced by the See also:action of the vibratile See also:cilia which are abundant along special tracts on the sides and roof of the See also:vestibule of the mouth and in the walls of the perforated pharynx ("ciliary ingestion"). Amphioxus favours a littoral See also:habitat, and rarely if ever descends below the 50-See also:fathom See also:line. Species occur in all seas of the temperate, tropical and subtropical zones. The See also:European species. A. lanceolatus, is found
in the See also:Black and Mediterranean Seas, and on the coasts of See also:France, Great See also:Britain and Scandinavia, while a closely allied species or subspecies, A. caribaeus, frequents the Caribbean region from Chesapeake to La See also:Plata. A. californiensis occurs on the coast of See also:California, and A. belcheri extends its See also:area of distribution from See also:Queensland through See also:Singapore to See also:Japan. A recently described species, Dolichorhynchus indicus, characterized by the great length of the praeoral See also:lobe or snout, has been dredged in the See also:Indian Ocean, Paramphioxus bassanus occurs on the coast of See also:Australia from See also:Port See also:Phillip to Port See also:Jackson; P. cingalensis at See also:Ceylon. Epigonichlhys cultellus (fig. I) inhabits Torres Strait, and has also been
body (these have no See also:constant numerical relation to the myomeres); M, notochord as seen through the transparent myotomes, the thin See also:double-lined spaces being the connective-See also:tissue septa and the broader spaces the muscular tissue of the myotomes; N, position of See also: Lettering as in (2). (¢) See also:Section taken transversely through praeoral region near termination of nerve tube. a, Olfactory ciliated See also:pit on animal's left side, its wall confluent with substance of nerve tube; b, pigment spot (rudimentary See also:eye) on anterior termination of nerve tube; c, first pair of nerves in section; d, fin See also:ray; e, myotome; f, notochord; g, space See also:round myotome (?artifact or coelom) ; h, subchordal canal (? See also:blood-vessel); i, a symmetrical epipleura of praeoral See also:hood. found at See also:Ternate. Asymmetron lucayanum is the Bahaman representative of the family, with a subspecies, A. caudatum, in the See also:South Pacific from New See also:Guinea to the See also:Loyalty Islands. The Peruvian species, Branchiostoma elongatum, with nearly eighty myotomes, cannot at See also:present be assigned to its proper subgenus. See also: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 See also:appearance, showing iridescent See also:play of See also:colour. The body is narrow, laterally compressed and pointed at both ends. The See also:main musculature can be seen through the thin skin to be divided into about sixty pairs of muscle-segments (myotomes) by means of See also:comma-shaped dissepiments, the myocommas, which stretch between the skin and the central skeletal See also:axis of the body. These myotomes enable it to swim rapidly with characteristic See also:serpentine undulations of the body, the movements being effected by the alternate contraction and relaxation of the See also:longitudinal muscles on both sides. Apparently correlated with this See also: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) See also:early undergo a certain distortion, the effect of which is to carry the somites of the left side forwards through the length of one See also:half-segment. For example, the twenty-seventh myotome of the left side is placed opposite to the twenty-See also:sixth myocomma of the right side. The back of the body is occupied by a'See also:crest, called the dorsal fin, consisting of a hollow See also:ridge, the cavity of which is divided into about 250 compartments or fin See also:chambers, into each of which, with the exception of those near the anterior and posterior end of the body, projects a stout See also:pillar composed of characteristic laminar tissue, the fin ray. The dorsal crest is continued round both extremities, becoming See also: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 See also: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 See also:supply and musculature by which the cirri can be either spread out, or See also:bent inwards so that those of one side may interdigitate with those of the other, thus completely closing the en-See also: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 See also:convex, so that the body presents the appearance of a spherical triangle in transverse section, the See also: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 See also:lymph-space, the metapleural canal. In the genus Branchiostoma the See also: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 See also: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-See also:system, namely, the ventral fin, which is composed of two portions, a See also:lower See also: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 See also: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 See also:middle line to the left side of the fin. In Asymmetron the caudal region is remarkable for the curious See also: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 See also:purification of the blood, they are not usually considered together, and it is therefore the more remarkable that their See also:close association in Amphioxus renders it necessary to treat them in See also: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 See also: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 See also:tongue-See also: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 See also:life of the animal. The gill-clefts open directly from the cavity of the pharynx into that of the See also:atrium, and so give See also: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 See also:separate and See also:divide the successive gill-clefts from one another are traversed by blood-vessels which run from a See also:simple tubular See also: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 See also:heart. As water for respiration streams through the clefts, gaseous interchange takes See also: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 See also: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 See also: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 See also:series of bags or pockets, which may be called the atrial pouches. At the See also:top of each of these pouches there is a See also: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 See also: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 See also: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 See also: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 See also:liver. Nervous System.—As has already been indicated, a solid sub-cylindrical elastic See also: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 See also:column. Immediately above the notochord there lies another elongated slit when relaxed (as in the See also: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 See also:blind pouch behind the atriopore as far as the anus. subcylindrical See also:cord, also surrounded by a sheath of connective tissue. This cord is neither elastic nor solid, but consists of nerve tissue, See also:fibres and ganglion cells, surrounding a small central canal. For the See also: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 See also:brain and See also: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 See also: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 See also: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- See also:mental in position. In addition to the cerebral eyespot there are large See also: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. See also:Hesse), each consisting of two cells, a See also:cup-shaped pigment See also: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 See also:illumination preferentially from the right side, although this fact has no relation with the side upon which Amphi- See also: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 See also: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 See also: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 (See also:figs. 2, 3, 4). At the breeding See also: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 See also: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 See also:pore; bc, rudiment of buccal See also:skeleton; c, cilia; cb, ciliated See also:band; cc, ciliated groove; cm, cilia at margin of mouth ; gl, external opening of See also: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; See also:rc, renal cells on atrial floor; rm, edge of right metapleur; so, sense See also:organ opening into praeoral pit ; ss, thickenings, the rudiments of the See also: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 See also: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 See also:hour after commencement of development the membrane ruptures and the See also:oval embryo escapes, See also:swimming by means of its flagella at the surface of the See also:sea for another twenty-four See also:hours, during which the principal organs are laid down, although the mouth does not open until the close of this See also: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 See also:bend up, meet, and finally fuse to form a tube, the medullary or neural tube. An important fact to See also: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 See also: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 See also: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 See also: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 See also:metamorphosis, which cannot be further described here. One point must not be omitted, namely, the homogeny of the endostyle of Amphioxus and the See also:thyroid gland of Craniata. and the Ancestry of the Vertebrates (1894) ; Remarks on some See also:recent See also:Work on the Protochorda," Quart. J. Micr. Sci. See also:xxiii. (1899), p. 223; " Dolichorhynchus indicus," ib. (1901); W. B. Benham, Heteropleuron of New See also:Zealand," ib. (1901) E Burchardt, " Finer See also:Anatomy of Amphioxus," with bibliography, See also:Jena Zeitschr. xxxiv. (1900), P. 719. (A. Additional information and CommentsThere are no comments yet for this article.
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