Alternation of Generations.—Fig. 66 represents an aggregated or sexual Salpa, which was once a member of a chain, since it shows a testis and a developing embryo. The ova (always few in number, usually only one) appear at a very early period in the developing chain Salpa, while it is still a part of the gemmiparous stolon in the body of the solitary Salpa. This gave rise to the view put forward first by Brooks that the ovary really belongs to the solitary stolon-bearing Salpa, which is therefore a female producing a series of males by asexual gemmation, and depositing in each of these an ovum, which will afterwards, when fertilised, develop in the body of the male into a solitary or female Salpa. This idea, if adopted, would profoundly modify our conception of Salpa as an example of a life-history showing alternation of generations, but it seems to me to give a distorted view of the sequence of events. The fact that the stolon while in the solitary Salpa contains, along with representatives of other important systems of the body, a row of germinal cells, does not constitute that solitary Salpa the parent of the ova which these germinal cells will afterwards become in the body of an independent bud. We must regard as the parent the body in which the ova become mature and fulfil their function. The sexual or chain Salpa, although really hermaphrodite in its life-history, is usually^[109] protogynous, i.e. the ova mature at an earlier period than the male organ or testis. This prevents self-fertilisation. The ovum is presumably fertilised by the spermatozoa of an older Salpa belonging to another chain, and the embryo is far advanced in its development before the testis is formed. The development takes place inside the body of the parent, and is "direct"—no tailed larval form being produced.

Development and Life-history.—The segmentation of the egg is holoblastic, and gives rise to a number of blastomeres, which are for a time masked by the phenomenal activity of certain cells of extraneous origin, the "kalymmocytes," derived from the follicular epithelium surrounding the ovum. These follicular kalymmocytes migrate into the ovum, surround groups of blastomeres, and arrange themselves so as to reproduce the essential structure of the future embryo for which they form what may be termed a scaffolding or temporary support. After a time the blastomeres become active, proliferate rapidly, and finally press upon and absorb the kalymmocytes, and so eventually take their proper place in building up the organs. Some observers regard the kalymmocytes as being passive and nutritive only in function.

At an early period in the development a part of the surface of the embryo, on its ventral edge, becomes separated off, along with a part of the wall of the cavity ("oviduct"—a diverticulum from the atrium) in which it lies, to form the "placenta" (Fig. 67, pl) in which the embryonic and maternal blood-streams circulate in close proximity, and so allow of the conveyance of nutriment to the developing embryo by means of large migrating placental cells. At a somewhat later stage a number of cells placed at the posterior end of the body alongside the future nucleus become filled up with oil-globules to form a mass of nutrient material—the "elaeoblast" (Fig. 67, ebl)—which is used up later in the development. Many suggestions have been made as to the homology and meaning of the elaeoblast; but it may now be regarded as most probable that it is reserve food-material associated with the disappearing rudiment of the tail found in the larval condition of most Ascidians. The development is direct; and it may be said, then, that this young asexual (solitary) Salpa differs from the corresponding form in the life-history of Doliolum (Fig. 60, A) in that its tail is no longer a locomotory organ, but is represented by a nutritive mass, the elaeoblast, while the body, in place of being free, is attached by its ventral surface to a special organ of nutrition—the "placenta"—in connexion with the blood-stream of the parent.

This embryo sexually produced inside the body of an aggregated form becomes a solitary Salpa (such as Fig. 61, B), which differs in appearance, structure, and habits from its parent, and has no reproductive organs. After swimming for a time, however, it develops the ventral stolon on which buds form which are eventually sexual Salpae. These are set free from the solitary form in sets, still connected together, and they may swim about together for a time as a chain of aggregated Salpae before separating to become the adult sexual individuals (such as Fig. 61, A).

Classification.—Salpa may be divided into the following subgenera:^[110]—Cyclosalpa, Blainville, in which the alimentary canal is ortho-enteric, and the "chain" consists of individuals united in a circle; Iasis, Savigny, with several embryos formed at a time; and Pegea, Sav., Thalia, Blumenbach, and Salpa, Forskål, all with one embryo only, and differing from one another in the condition of the "gill" and other details: all except Cyclosalpa have the alimentary canal caryo-enteric. Cyclosalpa has three species, the best known of which is C. pinnata of the Mediterranean, a form possessing light-producing organs like those of Pyrosoma, but placed along the sides of the body. Salpa has four or five species, one of which, S. runcinata-fusiformis (Fig. 61), has occasionally been found in British seas; Thalia includes the species T. democratica-mucronata, which has been sometimes obtained in swarms in the Hebridean seas, or cast ashore on our southern or western coasts; Pegea has the species P. scutigera-confoederata; and Iasis contains the remaining half-dozen species, the best known of which is I. cordiformis-zonaria, the only other Salpian which has been found in British seas.

The family Octacnemidae includes the single remarkable genus Octacnemus, now known in a solitary and an aggregated form. It was found during the "Challenger" expedition, and was first described by Moseley. It is apparently a deep-sea representative of the pelagic Salpidae, and may possibly be fixed at the bottom. The body in the solitary form is somewhat discoid, with its margin prolonged to form eight tapering processes, on to which the muscle-bands of the mantle are continued. The alimentary canal forms a compact nucleus, which is attached to an apparently imperforate membrane which stretches across the body, separating the branchial from the atrial cavities. The endostyle is very short, and the dorsal lamina is also much reduced. The reproduction and life-history are entirely unknown. The aggregated form consists of a small number of individuals united by a slender cord composed of test, body-wall, and endodermal tissue. Octacnemus has been found^[111] in the South Pacific from depths of 1070 and 2160 fathoms, and off the Patagonian coast from 1050 fathoms. Two species have been described: O. bythius, Moseley, and O. patagoniensis, Metcalf. Metcalf, who has recently investigated the aggregated form (O. patagoniensis), considers that the genus is more nearly related to the Clavelinidae than to the Salpidae. Possibly its position might be best indicated by a line diverging from near the point (3) in the phylogenetic diagram below.

General Conclusions.

The following diagram is a graphic representation of the genetic affinities, or what is now generally supposed to have been the probable course of phylogeny of the Tunicata. It will be noticed that it shows (1) the Proto-Tunicates arising from Proto-Chordata, not far from the ancestors of Amphioxus (see also, this vol. p. 112); (2) that the Larvacea are regarded as the most primitive section of the group; (3) that the Thaliacea (Doliolidae and Salpidae) are supposed to be derived not directly from primitive pelagic forms, but through the early fixed Ascidians, not far from (4) the ancestral compound Ascidians, which gave rise to the Pyrosomatidae; (5) that the Ascidiidae and other higher Simple Ascidians are derived, like the Compound Ascidians, from ancestral Clavelinidae; and (6), that the Ascidiae Compositae are polyphyletic, the Holosomata (Botryllidae and Polystyelidae) being derived from ancestral Simple Ascidians independently of the Merosomatous families.

The Tunicata are remarkable for the variety in appearance, structure, and life-history which they present. No group illustrates in a more instructive manner so large a number of important biological principles and phenomena. They show solitary and colonial forms, fixed and free, pelagic and abyssal. The development is in some cases larval and with metamorphosis, in others abbreviated and direct. Persistent traces of ancestral characters are seen in the embryonic and larval stages, while the adults present the most varied secondary adaptations to littoral, pelagic, and deep-sea, free-swimming and sessile modes of existence. In the details of their classification they demonstrate both stable and variable species, monophyletic and polyphyletic groups. They exhibit the phenomena of gemmation and of embryonic fission, of polymorphism, hibernation, alternation of generations, and change of function. They have long been known as a stock example of degeneration; but in fact they lend themselves admirably to the exposition of more than one "Chapter of Darwinism."

* * * * *

Note to P. 78.—Oligotrema, Bourne (Quart. J. Micr. Sci. xlvii. Pt. ii. 1903, p. 233), a Molgulid from the Loyalty Islands, has a reduced branchial sac and greatly developed pinnate, muscular branchial lobes, probably used in capturing food.

CHAPTER IV

CEPHALOCHORDATA

INTRODUCTION—GENERAL CHARACTERS—ANATOMY OF AMPHIOXUS—EMBRYOLOGY AND LIFE-HISTORY—CLASSIFICATION OF CEPHALOCHORDATA—SPECIES AND DISTRIBUTION

The Cephalochordata comprise only a small group of little fish-like forms, the Lancelets, usually known as "Amphioxus," and referable to about a dozen species arranged in several closely allied genera under the single family Branchiostomatidae. The best known form is Branchiostoma lanceolatum (Pallas), the common Amphioxus or Lancelet, which has been found in British seas, and even as far north as the coast of Norway, but is much more common in warmer waters, such as the Mediterranean, and is also found in the Indian Ocean. It is abundant in the Bay of Naples, and lives and breeds in great numbers in a salt lagoon, the "Pantano," near Messina, and from these localities most of the specimens have been obtained for the numerous recent researches upon its structure and development.

Amphioxus was first discovered and described (1778) by Pallas, who regarded it as a Mollusc, and named it Limax lanceolatus. It was first correctly diagnosed as a low Vertebrate, and named Branchiostoma, by Costa, in 1834. The term Amphioxus, under which it has become so well known, was applied to it a couple of years later by Yarrell.

The anatomy was for the first time fully investigated by Johannes Müller in 1841, and this important memoir has been supplemented in regard to special systems and histological details by numerous papers by many leading zoologists, such as those by Huxley in 1874, Langerhans in 1876, Lankester in 1875 and in 1889, Retzius in 1890, and Boveri and Hatschek, both in 1892. Important papers on special points have also been written by Rolph, Rohde, Benham, Andrews, Goodrich, and others. The development was first elucidated by Kowalevsky in 1867, at about the same time when he studied the development of the Ascidians, and later again in 1877. Further papers on the development and metamorphosis we owe to Hatschek in 1881, Lankester and Willey in 1890 and 1891, Wilson in 1893, and quite recently to MacBride. Dr. Willey's book, Amphioxus and the Ancestry of the Vertebrata (1894), contains a summary of investigations on structure and development, an interesting discussion of the relations of Amphioxus to the other Chordata, and a full bibliography.

In addition to such original researches, Amphioxus is studied in more or less detail every year by countless senior and junior students in zoological laboratories and marine stations throughout the civilised world. The value of this primitive form as an object of biological education depends upon the fact that it shows the essential Vertebrate characters, and their mode of formation, in a very simple and instructive condition. Although no doubt somewhat modified, and possibly degenerate in some details of structure, in its general morphology it presents us with a persistent type probably not far removed from the ancestral line of early Chordata. There are no sufficient grounds for the view that Amphioxus is a very degenerate representative of fish-like Vertebrata.

General Characters.—The Cephalochordata (or Acrania, in contradistinction to the Craniata or Vertebrata) are marine, non-colonial Chordata, in which the notochord extends the entire length of the body, running forward into the snout beyond the nervous system. There is no skull, and the notochord is not surrounded by any vertebral column. There are no limbs nor paired fins. There is no exoskeleton, and the ectoderm is a single layer of non-ciliated columnar cells. The mouth is ventral and anterior, the anus is ventral, posterior, and asymmetrically placed on the left side. The pharynx is a large branchial sac, having its sides perforated by many gill-slits, and is surrounded by an ectodermal enclosure, the atrium, which opens to the exterior by a median ventral atriopore. The stomach gives off a simple saccular pouch, the liver, which has connected with it a simple hepatic portal blood system. There is a respiratory circulation, the contractile ventral vessel which represents the heart sending the colourless blood forward to the respiratory pharynx to be purified. The body-wall is segmented into over fifty myotomes. There are numerous separate nephridia which develop from the mesoderm and open into the atrium. The brain remains undeveloped, being scarcely distinct from the spinal cord. There are two pairs of cerebral nerves, and many spinal, in which the dorsal and ventral roots or nerves do not unite. The sense-organs are simple; there are no paired eyes and no auditory organs. The sexes are separate; the gonads are metamerically arranged on the body-wall, and have no ducts: they burst into the atrium. In the development the segmentation is complete, a gastrula is formed by invagination, the nervous system is formed from the dorsal epiblast, the notochord from the hypoblast, and the mesoderm arises from metameric coelomic pouches. The body-cavity is an enterocoele. The gill-slits are at first perforations of the body-wall opening from the pharynx to the exterior, which later become enclosed by the development of the atrium.

Anatomy.

External Characters.—Amphioxus^[112] is about 1½ to 2½ inches in length, slender, somewhat translucent, and pointed at both ends (Fig. 69). It lives in shallow water and burrows in the sand, head first, with great rapidity. It frequently remains with the anterior end protruding from the sand. When on the surface it lies on one side. It is said to swim freely at night. The head end is rather the thicker, and the anterior two-thirds of the ventral surface are flattened (Fig. 70, A), and may be slightly ridged longitudinally. The lateral edges of this flat area project as metapleural folds (Fig. 70, mt.pl), which begin anteriorly at the edges of the external mouth, and die away in the middle line posteriorly behind a median opening, the atriopore (Fig. 70, atrp). From this point a ventral median fin (vent.f) extends backwards around the pointed posterior end (caudal fin, cd.f), and then forwards along the upper surface (dorsal fin, dors.f) to the anterior end of the body. These fins thus constitute a continuous median fold around a great part of the animal (Fig. 70, B, and Fig. 71).

The surface is soft all over, there being no exoskeleton. The epidermis or ectoderm is formed by a single layer of epithelial cells (see Fig. 72, p. 118), some of which bear sensory processes, while others have a striated cuticular border. There is no general ciliation of the surface in the adult.

The true mouth is a small pore at the bottom of a large vestibule (the stomodaeum), placed at the anterior end of the ventral surface (Figs. 70 and 71), and formed by the "oral hood," which may be a prolongation forwards of the atrial or metapleural folds at each side. The edges of the oral hood bear 12 to 20 pairs of cirri (Fig. 70, cir) or ciliated tentacles (strengthened by skeletal rods), which form a sensory fringe around the opening. The anus (Figs. 70 and 71, an), is asymmetrical, being placed on the left side of the ventral fin, some distance behind the atriopore, and not far from the posterior end of the body. The short region behind the anus and surrounded by the caudal fin may properly be called "tail." The current of water for respiratory and nutritive purposes, and which may carry the ova and spermatozoa to the exterior, usually passes in at the mouth and out at the atriopore, as in the Tunicata. On occasions, however, it is said to be reversed.

General Structure.—The general plan of organisation of the body (see Fig. 71) is that a longitudinal skeletal axis, the notochord (nch), separates a dorsal nervous system (sp.cd) from a ventral reduced coelom (coel), in which lie the alimentary canal (int), the gonads (gon), and other organs. Thus a transverse section of the body (see Fig. 72) shows the typical Chordate arrangement of parts, and is comparable with a transverse section of a tadpole, a young fish, or a larval Ascidian. A peribranchial (atr) or atrial cavity (which is morphologically a part of the external world shut in) lies external to the coelom and body-wall around the pharynx and the greater part of the alimentary canal, and opens to the exterior by the atriopore. As in the Tunicata, the perforations (gill-slits) in the wall of the pharynx (br.cl) open into the atrial cavity and so indirectly to the exterior.

Musculature.—The thick body-wall is largely formed by muscular tissue metamerically segmented into about 60 myotomes (Fig. 71, myom). These muscle-masses, which (as is usual in Vertebrata) are thickest dorsally at the sides of the notochord and spinal chord (Fig. 72, m), are so arranged as to present the appearance in a lateral view of the body of a series of shallow cones (<<) fitting into one another and with their apices directed forwards. The muscle fibres are striated, and run longitudinally along the body from the anterior to the posterior edge of each myotome, so as to be attached at their ends to the two septa of connective tissue which form the boundaries of the myotomes. These septa, the myocommas, are conspicuous features in the external appearance of the body (Fig. 70, B). They are not arranged so as to be opposite one another on the two sides, but the myotomes on the right and left sides alternate, as can be seen in a transverse section (Fig. 74, A, p. 121).

It is by means of these lateral muscle-bundles that the rapid vibration or alternate bending of the body from side to side in swimming or burrowing can be performed. There are usually, on each side, 35 myotomes in front of the atriopore, 14 between the atriopore and the anus, and 11 postanal, making 60 in all: some species have only about 50 myotomes, and some as many as 85. (See Classification, p. 137, where a list of the species with the number of myotomes in each is given.)

There are also transverse muscles (Fig. 72, mt) extending across the ventral surface in the region of the body enclosed by the metapleural folds, and serving to compress the atrial cavity, and so aid in the expulsion of its contents.

Outside the muscular layer of the body-wall the thin integument is formed of a dermal layer of soft connective tissue, covered by the epidermis, a single layer of columnar cells, many of which, especially on the oral cirri, have sensory bristles.

Skeleton.—The endoskeleton consists of the notochord and some tracts of modified connective tissue which support various parts of the body.

The notochord of this animal is noteworthy amongst Chordata for extending practically the entire length of the body, including the head, from snout to tip of tail (Fig. 71). It lies in the median plane, but nearer the dorsal than the ventral surface (Fig. 72), and has the myotomes at its sides, the nervous system above and the alimentary canal below. It is elliptical in section, and tapers to the two ends. The nuclei of the original notochordal cells are displaced to the dorsal and ventral edges, and the greater parts of the cells, in the adult, are occupied by large vacuoles filled with a fluid secretion, so as to form by their distended condition a stiff elastic structure. This state of the cells, and the appearance it gives rise to (Fig. 73), seen best in young specimens, is very characteristic of notochordal tissue. Around the notochord lies a sheath of connective tissue which is continuous with the similar sheath around the nervous system and with the septa between the myotomes.

In addition to these skeletal layers of connective tissue there is a cartilage-like tract in the oral hood. This is jointed, or made up of separate rod-like pieces, one at the base of each cirrus, into which it sends a prolongation (Fig. 71, sk). The dorsal and ventral fins are supported by single and double rows respectively of what have been called "fin-rays." They are short rods of gelatinous connective tissue, each enclosed in a lymph space. Finally, the bars constituting the walls of the pharynx between the gill-slits contain slender skeletal rods which run obliquely dorso-ventrally, and are of a stiff, gelatinous nature (see Fig. 75, p. 122). This skeletal connective tissue consists in all cases of a fibrous deposit or matrix produced by the layer of epithelium (ectodermal, endodermal, or mesodermal) which adjoins the tissue.

Alimentary Canal.—This has, as its most noteworthy feature, the Chordate characteristic that the pharynx gives rise to the respiratory organ (see Figs. 71 and 74, A); and in size and prominence, both in side view and in sections, the modified pharynx of Amphioxus is fairly comparable with the branchial sac (pharynx) of many Tunicata (see Fig. 23, p. 51), and might be called by the same name.

The small primitive mouth, at the bottom of the cavity bounded by the oral hood (stomodaeum), has a membranous border, the velum (Fig. 71, vl), the edges of which are prolonged into a circle of 10 or 12 (up to 16 in some species) simple oral tentacles turned inwards towards the pharynx (compare tentacles of Ascidians, p. 45).

The pharynx, by far the largest part of the alimentary canal, and extending nearly half-way along the body, is more important as a respiratory than as a nutritive organ. Its walls over nearly the whole extent are perforated by a large, and indefinite, number (100 or more on each side) of gill-slits which run on the whole dorso-ventrally, but in the contracted condition seen in preserved specimens have their lower ends directed obliquely backwards, so that a vertical transverse section may cut through a number of such slits and the intervening branchial bars (Fig. 74, A, kb). These bars, and therefore the slits between them, are of two orders, primary and secondary, the latter being developed later in larval life as downgrowths or "tongue-bars," one from the top of each primary gill-slit, so as to divide it into two secondaries. The primary and the secondary (or tongue-) bars can be distinguished from one another by their structure in the adult animal (Fig. 75, A and B).

It must be remembered that these branchial bars, or septa between the gill-slits, are not merely portions of the wall of the pharynx, but are in a sense portions of the body-wall as well, and correspond in nature, though not in number, to the visceral arches in a Vertebrate lying between the visceral clefts which open on the exterior. In the adult Amphioxus the clefts in the wall of the pharynx do not open directly to the exterior, but into the peribranchial cavity or atrium, which, however, is only formed at a late larval period as an invagination or enclosure of ectoderm. Previous to that the first formed gill-slits opened to the exterior in Amphioxus (see larva, Fig. 86, p. 134), just as they do in a fish or a young tadpole. The atrial cavity is therefore, from its origin, lined by ectoderm, and the outer surface of a branchial bar is virtually a part of the outer surface of the body. It is only natural then to find that each bar contains a small section of the coelom in its interior, communicating dorsally and ventrally with other parts of that cavity (see Figs. 75 and 76). There are also blood-vessels which run in the branchial bars and their junctions. The greater part of the epithelium covering a branchial bar is pharyngeal epithelium or endoderm (Fig. 75, br.ep), but the external, wider, non-ciliated cells (Fig. 75, at.ep) are ectodermal cells lining the atrium. The gelatinous skeletal rods in the primary bars are forked ventrally, while those in the secondary bars are simple; and there are other points of detail in which the two kinds of bar differ. These bars are obviously more numerous in the adult than the myotomes, but in the young larva the first formed gill-clefts are metamerically arranged, and then later they increase greatly in number. It is the cilia covering the pharyngeal epithelium on the branchial bars, possibly aided by the ciliated tracts of the oral hood, which cause the current of water already alluded to.

Transverse branchial junctions (synapticula) run across the branchial bars, connecting them at frequent intervals, and these transverse connexions, like the branchial bars, are supported by skeletal rods. Along the ventral median line of the pharynx runs a groove, the endostyle or hypopharyngeal groove, comparable with the similar structure in the branchial sac of Tunicata. This longitudinal groove (Fig. 76, gl) is lined by ciliated epithelium containing four tracts of gland cells (compare endostyle in Ascidians, Fig. 20, p. 46). There is reason to believe that this organ is the homologue of the thyroid gland of Vertebrata. As in the case of Tunicata the endostyle secretes mucus, which is carried forwards by the cilia to constitute a train with entangled food particles which pass back dorsally to the stomach. At the anterior end the ciliated lips of the endostyle diverge to the right and left to encircle the front of the pharynx as the peripharyngeal bands. These unite again dorsally to form the epipharyngeal (or hyperpharyngeal) groove which leads backwards, corresponding to the hypopharyngeal groove below (see Fig. 74, A), till the posterior end of the pharynx is reached.

The remainder of the simple alimentary canal is straight, and is scarcely differentiated into regions. A slight narrowing of the tube behind the pharynx has been called the oesophagus, and a slight enlargement which follows, the stomach. From this point the intestine tapers backwards to the anus (Fig. 71, p. 116). The ventral edge of the stomach gives off a blind pouch, the hepatic caecum or saccular liver, which runs forwards on the right-hand side of the pharynx (Fig. 74, A, l). This is a digestive gland, is lined with glandular epithelium, and apparently corresponds with the liver of Vertebrata. There are no other digestive glands in connexion with the alimentary canal of Amphioxus.

Coelom.—In the young larva there are at first (as in Balanoglossus) five coelomic spaces, a median anterior "head-cavity," a pair of antero-lateral "collar-cavities," and a pair of more posterior long lateral grooves from which arise, in the later larva, the segmented myotomes and ventrally a large coelomic space surrounding the alimentary canal and separating it from the body-wall. In the adult animal, however, the coelom has been so much displaced by the formation of the spacious atrium that in front of the atriopore it can only be recognised as a series of canals and crevices. The relations of coelom to atrium in the region of the intestine are seen in Fig. 74, B, and in the region of the pharynx in Fig. 74, A. Fig. 72 shows the distribution of the spaces more in detail (see also Fig. 71). Beginning anteriorly, along the dorsal surface of the pharynx and beneath the notochord run a pair of dorsal coelomic canals, one at each side of the epipharyngeal groove; these give off ventral diverticula which pass down the primary branchial bars of the pharyngeal wall and unite ventrally in a median tube, the endostylar coelom (see Fig. 72, ec). At the posterior end of the pharynx these dorsal and ventral canals unite in a narrow coelomic space encircling the stomach, inside the wall of the atrium, and sending an extension forwards around the liver (Fig. 74, A, l). In the region of the intestine, behind the atriopore, the coelom is allowed to expand to its primitive condition on the left-hand side (Fig. 74, B), but is still reduced on the right side, where there is a prolongation of the atrial cavity reaching nearly to the anus. All these coelomic spaces are lined by a coelomic epithelium.

The Blood System of Amphioxus, although as simple as that of a Chaetopod worm, is undoubtedly laid down on the Vertebrate plan—even though there is no distinct heart and the vessels are few and of simple structure. Capillary networks are formed in some places, but the colourless blood also extends into many lacunae or lymph spaces, such as those around the fin-rays and in the metapleura. As in a typical lower Vertebrate, there is a contractile ventral vessel (the ventral or branchial aorta, Fig. 77, v.ao) running forwards under the alimentary canal to the pharynx, and giving off on each side afferent branchial vessels, which pass up the primary branchial bars and give off branches joining the vessels in the secondary bars. These latter do not communicate directly with the ventral aorta, but the vessels in all the branchial bars open dorsally by efferent branchial vessels into the paired dorsal aortae (Fig. 77, d.ao), which run backwards along the top of the pharynx, one at each side of the epipharyngeal groove. In the vessels of the branchial bars and their connectives the blood is aerated by the current of water passing through the gill-slits, and so reaches the dorsal aortae in a purified condition. The right-hand dorsal aorta is continued forward further into the snout than its fellow of the other side, and is dilated at its extremity (Fig. 77). At the posterior end of the pharynx the paired dorsal aortae unite to form the median dorsal aorta which runs backwards, lying between notochord and alimentary canal. This vessel gives off branches to the wall of the intestine, and these break up into capillary networks (Fig. 77, cp), from which the blood is collected by the median sub-intestinal vein. This then flows forwards to pass by the hepatic portal vein to the ventral edge of the saccular liver, in the wall of which it is distributed in a capillary network. The blood is collected on the dorsal edge of the liver by the hepatic vein, which runs posteriorly and then turns downwards and forwards to become continuous with the posterior end of the ventral aorta or "heart."

It is clear that this course of the circulation agrees with that of a typical lower Vertebrate in all essential points:—(1) in having the main artery a dorsal aorta in which the blood flows backwards; (2) in having a ventral vessel representing the heart, and sending impure blood forwards to the respiratory region of the alimentary canal to be aerated; and (3) in having a hepatic portal system consisting of the capillaries of the liver, through which the blood from the intestinal wall has to pass before reaching the ventral vessel (heart).

Renal Excretory functions have been attributed to various organs in Amphioxus, and it is quite possible that, in addition to the true nephridia which are now known, other tracts of tissue in the body may be able to eliminate nitrogenous waste matters. Such are certain clumps of columnar epithelial cells on the floor of the atrium, and the single pair of large brown atrio-coelomic funnels lying on the dorsal edge of the posterior end of the pharynx (Fig. 71, br.f). There are, however, a large number (about 100 pairs) of minute nephridia, discovered (1890) by Weiss and by Boveri independently, lying at the sides of the dorsal coelomic canals above the pharynx, which must be regarded as the chief functional renal organs. These are bent tubules, partly glandular and partly ciliated, each giving off several caecal knobs (at first supposed to be open nephrostomes, one shown at each end of the tubule and three along its upper surface in Fig. 78), which project into the coelom, and opening by one nephridiopore (on the lower surface, and opposite a tongue bar of the pharynx) into the atrial cavity. The knobs, or closed nephrostomes, are surrounded by peculiar, slender, club-shaped tubular and flagellated cells—which Goodrich^[113] has shown to correspond to the "solenocytes" in the nephridia of Polychaete worms (see Fig. 79).