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Fig. 1.—Lineus geniculatus. (From Bürger.) 1, Lateral slits on head; 2, anus. |
NEMERTINA, or Nemerteans (Nemertea), a subdivision of worms,[1] characterized by the ciliation of the skin, the presence of a retractile proboscis, the simple arrangement of the generative apparatus, and in certain cases by a peculiar pelagic larval stage to which the name “pilidium” has been given. Many of them are long thread-shaped or ribbon-shaped animals, more or less cylindrical in transverse section, Even the comparatively shortest species and genera can always be termed elongate, the broadest and shortest of all being the parasitic Malacobdella and the pelagic Pelagonemertes. There are no exterior appendages of any kind. The colours are often very bright and varied. Nemertines live in the sea, some being common amongst the corals and algae, others hiding in the muddy or sandy bottom, and secreting gelatinous tubes which ensheath the body along its whole length. Formerly, they were generally arranged amongst the Platyelminthes as a sub-order in the order of the Turbellarians, but with the advance of our knowledge of these lower worms it has been found desirable to separate them from the Turbellarians and to look upon the Nemertina as a separate phylum.
O. Bürger classifies Nemertines into four orders:—
I. Protonemertini, in which there are two layers of dermal muscles, external circular and internal longitudinal; the nervous system lies external to the circular muscles; the mouth lies behind the level of the brain; the proboscis has no stylet; there is no caecum to the intestine. Families, Carinellidae, Hubrechtiidae.
II. Mesonemertini, in which the nervous system has passed into the dermal muscles and lies amongst them; other characters as in Protonemertini. Family, Cephalothricidae.
III. Metanemertini, in which the nervous system lies inside the dermal muscles in the parenchyma; the mouth lies in front of the level of the brain; the proboscis as a rule bears stylets; the intestine nearly always has a caecum. Families, Eunemertidae, Ototyphlonemertidae, Prosorhocmidae, Amphiporidae, Tetrastemmatidae, Nectonemertidae, Pelagonemertidae, Malacobdellidae.
This order represents the Hoplonemertini of Hubrecht.
IV. Heteronemertini, in which the dermal musculature is in three layers, an external longitudinal, a middle circular, an internal longitudinal; the nervous system lies between the first and second of these layers; the outer layer of longitudinal muscles is a new development; there is no intestinal caecum; no stylets on the proboscis and the mouth is behind the level of the brain. Families, Eupolidae, Lineidae.
This order represents the Schizonemertini of Hubrecht and the family Eupolidae.
The first three orders, which have a double muscular layer, external circular and internal longitudinal, are sometimes grouped together as the Dimyaria; the Heteronemertini, in which a third coat of longitudinal muscles arises outside the circular layer, are then placed in a second branch, the Trimyaria.
The following families and genera are represented on the British coasts: Carinellidae, Carinella; Cephalothricidae, Cephalothrix, Carinoma; Eunemertidae, Eunemertes; Ototyphlonemertidae, Ototyphlonemertes; Amphiporidae, Amphiporus, Drepanophorus; Tetrastemmidae, Tetrastemma, Prosorhocmus; Malacobdellidae, Malacobdella; Eupoliidae, Eupolia, Valencinia, Oxypolia; Lineidae, Lineus, Euborlasia, Micrura, Cerebratulus, Micrella.
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Fig. 2.—Diagrams of the organs of a Nemertine. A, From below; B, from above. | ||||
| m, | Mouth. | Br, | Brain-lobes. | |
| div, | Intestinal diverticula. | ln, | Longitudinal nerve stems. | |
| a, | Anus. | pr, | Proboscis. | |
| ov, | Ovaries. | ps, | Proboscidian sheath. | |
| n, | Nephridia. | p.o., | Opening for proboscis. | |
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Fig. 3.—Anterior portion of the body of a Nemertine. | |
| Br, | Brain-lobes. |
| N, | Lateral nerves. |
| PS, | Proboscidian sheath. |
| Pr, | Proboscis. |
| P.o., | Exterior opening through which the proboscis is everted or rhynchostome. Oesophagus and mouth shown by dotted lines. |
Anatomy.—Proboscis and Proboscidian Sheath.—The organ most characteristic of a Nemertine is without doubt the proboscis. With very few exceptions (Malacobdella, Akrostomum, where it has fused with the mouth into a single exterior opening), there is a terminal opening, the rhynchostome (subterminal in Valencinia), at the foremost tip of the body, out of which the proboscis is seen shooting backwards and forwards, sometimes with so much force that both its interior attachments are severed and it is entirely expelled from the body. It then often retains its vitality for a long time, apparently crawling as if it were itself a worm, a phenomenon which is at least partially explained by the extraordinary development of nervous tissue, equally distributed all through the walls of the proboscis, and either united into numerous longitudinal nerve-stems (Drepanophorus, Amphiporus) or spread out into a uniform and comparatively thick layer (Cerebratulus, sp.). This very effective and elaborate innervation, which has been directly traced to the brain, whence strong nerves (generally two) enter the proboscis, renders it exceedingly probable that the most important functions of the proboscis are of a sensiferous, tactile nature. In Nemertines the everted proboscis is retracted in the same way as the tip of a glove finger would be if it were pulled backwards by a thread situated in the axis and attached to the tip. The comparison may be carried still further. The central thread just alluded to is represented in the Nemertean proboscis by that portion which is never everted, and the tip of the glove by the boundary between the evertible and non-evertible portion of the proboscis—a boundary which in the Metanemertini is marked by the presence of a pointed or serrated stylet. This stylet is thus situated terminally when the proboscis has reached its maximum eversion. It adds a decisively aggressive character to an organ the original significance of which, as we have seen, was tactile. This aggressive character has a different aspect in several genera which are destitute of a central stylet, but in which the surface that is turned outwards upon eversion of the proboscis is largely provided with nematocysts, sending the urticating rods of different sizes in all directions. In others this surface is beset with thick, glandular, adhesive papillae.
The comparison with the glove-finger
is in so far insufficient as the greater
portion of the non-evertible half of the
proboscis is also hollow and clothed by
glandular walls. Only at the very hindermost
end does it pass into the so-called
retractor-muscle (fig. 2), which is
attached to the wall of the space, or
rhynchocoel, in which the proboscis moves
about. This retractor-muscle, indeed,
serves to pull back with great rapidity,
the extruded proboscis, and is aided
in its action by the musculature of the
head. The extrusion itself depends
entirely upon contraction of the muscular
walls of the space just mentioned, the
rhynchocoel. As it is (1) closed on all
sides, and (2) filled with a corpuscular
fluid, the contractions alluded to send
this fluid to impinge against the anterior
portion, where the proboscis, floating in
its sheath, is attached with it to the
muscular tissue of the head (fig. 3).
Partial extrusion lessening the resistance
in this region inevitably follows,
and when further contractions of the
walls of the sheath ensue total
extrusion is the consequence. It is
worthy of notice that in those Nemertines which make a very free
use of their proboscis, and in which it is seen to be continually
protruded and retracted, the walls of the proboscidian sheath are
enormously muscular. On the other hand, they are much less considerably
or even insignificantly so in the genera that are known
to make a rather sparing use of
their proboscis. The
rhynchocoel is formed by a split
which appears in the
mesoblast surrounding the
epiblastic pit which is the
forerunner of the proboscis. It
does not seem to be coelomic.
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| Fig. 4. | Fig. 5. | |
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Figs. 4, 5.—Proboscis with stylet, “reserve” sacs and muscular bulb of a Hoplonemertine. Fig. 4 retracted; fig. 5 everted. | ||
The proboscis, which is thus an eminently muscular organ, is composed of two or three, sometimes powerful, layers of muscles—one of longitudinal and one or two of circular fibres. In the posterior retractor the longitudinal fibres become united into one bundle, which, as noticed above, is inserted in the wall of the sheath. At the circular insertion of the proboscis in front of the brain the muscular fibres belonging to the anterior extremity of the body and those connected with the proboscis are very intimately interwoven, forming a strong attachment. The short tube between this circular insertion and the rhynchostome is called the rhynchodaeum.
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Fig. 6.—The armature from the proboscis of Drepanophorus. |
The proboscis broken off and expelled is generally reproduced, the posterior ribbon-like end of this reproduced portion again fusing with the walls of the sheath. There is reason to suppose that, when a wound is inflicted by the central stylet, it is envenomed by the fluid secreted in the posterior proboscidian region being at the same time expelled. A reservoir, a duct and a muscular bulb in the region (fig. 4) where the stylet is attached serve for this purpose. The significance of two or more (in Drepanophorus very numerous) small sacs containing so-called “reserve” stylets resembling in shape that of the central dart is insufficiently known.
The muscular walls of the rhynchocoel, which by their transverse contractions serve to bring about eversion of the proboscis in the way above traced, are attached to the musculature of the head just in front of the ganglionic commissures (fig. 3). In nearly all Nemertines the rhynchocoel extends backwards as far as the posterior extremity, just above the anus; in Carinella it is limited to the anterior body-region. The corpuscles floating in the fluid it contains are of definite shape, and in Cerebratulus urticans they are deep red, possibly from the presence of haemoglobin. They are usually larger than the blood corpuscles. Internally the muscular layers are lined by an epithelium. In the posterior portion this epithelium in certain Heteronemertea has a more glandular appearance, and sometimes the interior cavity is obliterated by cell-proliferation in this region. Superiorly the sheath either closely adheres to the muscular bodywall, with which it may even be partly interwoven, or it hangs freely in the connective tissue which fills the space between the intestine and the muscular body-wall.
Cutaneous System.—Externally in all species a layer of ciliated cells forms the outer investment. In it are, moreover, enclosed unicellular glands pouring their highly refracting contents, of a more or less rod-like shape, directly to the exterior. They appear to be the principal source of the mucus these animals secrete. In most Heteronemertines these elements are separated by a thin homogeneous basement membrane (fig. 8) from the following—that is, from a layer in which longitudinal muscular fibres are largely intermixed with tortuous glands, which by reason of their deeper situation communicate with the exterior by a much longer and generally very narrow duct. The pigment is also principally localized in this layer, although sometimes it is present even deeper down within the musculature. The passage from this tegumentary layer to the subjacent longitudinal muscular one is gradual, no membrane separating them. In Carinella, Cephalothrix, Polia and the Metanemertines the two tegumentary layers with their different glandular elements are fused into one; a thick layer of connective tissue is situated beneath them (instead of between them) and keeps the entire cutaneous system more definitely separate from the muscular (figs. 7, 8).
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| Fig. 7. | Fig. 8. | Fig. 9. | ||
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Figs. 7–9.—The layers of the body-wall in Carinella (fig. 7), the Metanemertini (fig. 8) and the Heteronemertini (fig. 9). c, Cellular tissue of the integument; Bm, basement membrane; circ. 1, outer circular, and long., longitudinal layer of muscular tissue; circ. 2, long. 1, additional circular and longitudinal layers of the same; nl, nervous layer. | ||||
Musculature and Connective Tissue.—The muscular layers by which the body-wall is constituted have been very differently and to some extent confusingly described by the successive authors on Nemertean anatomy. There is sufficient reason for this confusion. The fact is that not only have the larger subdivisions a different arrangement and even number of the muscular layers, but even within the same genus, nay, in the same species, well-marked differences occur. Increase in size appears sometimes to be accompanied by the development of a new layer of fibres, whereas a difference in the method of preparation may give to a layer which appeared homogeneous in one specimen a decidedly fibrous aspect in another. Nevertheless there are three principal types under which the different modifications can be arranged. One of them is found in the two most primitively organized genera, Carinella and Cephalothrix, i.e. an outer circular, a longitudinal and an inner circular layer of muscular fibres (fig. 7). The second is common to all the Heteronemertines, as well as to Polia and Valencinia, and also comprehends three layers, of which, however, two are longitudinal, viz. the external and the internal one, there being a strong circular layer between them (fig. 9). To the third type all the Metanemertini correspond; their muscular layers are only two, an external circular and an internal longitudinal one (fig. 8).
The Heteronemertini thus appear to have developed an extra layer of longitudinal fibres internally to those which they inherited from more primitive ancestors, whereas the Metanemertini are no longer in possession of the internal circular layer, but have on the contrary largely developed the external circular one, which has dwindled away in the Heteronemertini. The situation of the lateral nerve-stems in the different genera with respect to the muscular layers lends definite support to the interpretation of their homologies here given and forms the basis of Bürger's classification.
In Carinella, Cephalothrix and Polia, as well as in all Metanemertines, the basement membrane of the skin already alluded to is particularly strong and immediately applied upon the muscular layers. In the Heteronemertines there is a layer in which the cutaneous elements are largely represented below the thin basement membrane (fig. 8), between it and the bulk of the outer longitudinal muscles. The difference in the appearance of the basement membrane—sometimes wholly homogeneous, sometimes eminently fibrillar—can more especially be observed in differently preserved specimens of the genus Polia.
The connective tissue of the integument and basement membrane imperceptibly merges into that which surrounds the muscular bundles as they are united into denser and definite layers, and this is especially marked in those forms (Akrostomum) where the density of the muscular body-wall has considerably diminished, and the connective tissue has thus become much more prominent. It can then at the same time be observed, too, that the compact mass of connective tissue (“reticulum,” Barrois) which lies between the muscular body-wall and the intestine is directly continuous with that in which the muscular layers are embedded. Nuclei are everywhere present. The omnipresence of this connective tissue tends to exclude the formation of any perivisceral body cavity in Nemertines.
In Polia the connective tissue enclosed in the external muscular layer is eminently vacuolar—all the intermediate stages between such cells in which the vacuole predominates and the nucleus is peripheral and those in which the granular protoplasm still entirely fills them being moreover present.
In addition to the musculature of the proboscis and proboscidian sheath, longitudinal muscular fibres are found in the walls of the oesophagus, whilst transverse ones are numerous and united into vertical dissepiments between the successive intestinal caeca, thus bringing about a very regular internal metamerization. The genital products develop in intermediate spaces similarly limited by these dissepiments and alternating with the digestive caeca.
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Figs. 12.—The brain of a Nemertine, with its lobes and commissures. | ||||||||||
| S.N., | Nerves to sensory apparatus. | |||||||||
| P.N., | Nerves for proboscis. | |||||||||
| vag, | Nerves for oesophagus. | |||||||||
| L.N., | Lateral nerve-stems. | |||||||||
Nervous System and Sense Organs.—The nervous system of Nemertines presents several interesting peculiarities. As central organs we have to note the brain-lobes and the longitudinal lateral cords which form one continuous unsegmented mass of fibrous and cellular nerve-tissue. The fibrous nerve-tissue is more dense in the higher differentiated, more loose and spongy in the lower organized forms; the cellular nerve-tissue is similarly less compact in the forms that are at the base of the scale. No ganglionic swellings whatever occur in the course of the longitudinal cords. The brain must be looked upon as the anterior thickening of these cords, and at the same time as the spot where the two halves of the central nerve system intercommunicate. This is brought about by a double commissure, of which the ventral portion is considerably thicker than the dorsal, and which, together with the brain-lobes, constitutes a ring through which both proboscis and proboscidian sheath pass. The brain-lobes are generally four in number, a ventral and a dorsal pair, respectively united together by the above-mentioned commissures, and moreover anteriorly interfusing with each other, right and left. In Carinella this separation into lobes of the anterior thickenings of the cords has not yet commenced, the ventral commissure at the same time being extremely bulky. There is great probability that the central stems, together with the brain, must be looked upon as local longitudinal accumulations of nervous tissue in what was in more primitive ancestors a less highly differentiated nervous plexus, situated in the body-wall in a similar way to that which still is found in the less highly organized coelenterates. Such a nervous plexus indeed occurs in the body-wall of all Heteronemertines, sometimes even as a comparatively thick layer, situated, as are the nerve stems, between the external longitudinal and the circular muscles (fig. 9). In Carinella, where the longitudinal nerve-stems are situated exteriorly to the muscular layers, this plexus, although present, is much less dense, and can more fitly be compared to a network with wide meshes. In both cases it can be shown to be in immediate continuity with the coating of nerve-cells forming part of the longitudinal cords. It stretches forward as far as the brain, and in Carinella is again continued in front of it, whereas in the Heteronemertines the innervation of the anterior extremity of the head, in front of the brain, takes the form of more definite and less numerous branching stems. The presence of this plexus in connexion with the central stems, sending out nervous filaments amongst the muscles, explains the absence, in Pro-, Meso- and Heteronemertines, of separate and distinct peripheral nerve stems springing from the central stems innervating the different organs and body-regions, the only exceptions being the nerves for the proboscis, those for the sense organs in the head and the strong nerve pair (n. vagus) for the oesophagus. At the same time it renders more intelligible the extreme sensitiveness of the body-wall of the Nemertines, a local and instantaneous irritation often resulting in spasmodic rupture of the animal at the point touched.
In the Metanemertini, where the longitudinal stems lie inside the muscular body-wall, definite and metamerically placed nerve branches spring from them and divide dichotomously in the different tissues they innervate. A definite plexus can here no longer be traced. In certain Metanemertines the lateral stems have been noticed to unite posteriorly by a terminal commissure, situated above the anus, the whole of the central nervous system being in this way virtually situated above the intestine. In others there is an approximation of the lateral stems towards the median ventral line (Drepanophorus); in a genus of Heteronemertines (Langia), on the other hand, an arrangement occurs by which the longitudinal stems are no longer lateral, but have more or less approached each other dorsally.
In addition to the nerves starting from the brain-lobes just now especially mentioned, there is a double apparatus which can hardly be treated of in conjunction with the sense organs, because its sensory functions have not been sufficiently made out, and which will therefore rather be considered along with the brain and central nervous system. This apparatus is usually known under the name of the lateral organs. To it belong (a) superficial grooves or deeper slits situated on the integument near the tip of the head, (b) nerve lobes in immediate connexion with the nervous tissue of the brain, and (c) ciliated ducts penetrating into the latter and communicating with the former. Embryology shows that originally these different parts are separately started, and only ultimately become united into one. Two lateral outgrowths of the foremost portion of the oesophagus, afterwards becoming constricted off, as well as two ingrowths from the epiblast, contribute towards its formation, at least as far as both Meta- and Heteronemertines are concerned. As to the Mesonemertini, in the most primitive genus, Carinella, we do not find any lateral organs answering to the description above given. What we do find is a slight transverse furrow on each side of the head, close to the tip, but the most careful examination of sections made through the tissues of the head and brain shows the absence of any further apparatus comparable to that described above. Only in one species, Carinella inexpectata, a step in advance has been made, in so far as in connexion with the furrow just mentioned, which is here also somewhat more complicated in its arrangement, a ciliated tube leads into the brain, there to end blindly amidst the nerve-cells. No other intermediate stages have as yet been noticed between this arrangement and that of the Heteronemertini, in which a separate posterior brain-lobe receives a similar ciliated canal, and in which the oesophageal outgrowths have made their appearance and are coalesced with the nerve-tissue in the organ of the adult animal. The histological elements of this portion remain distinct both by transmitted light and in actual sections.
These posterior brain-lobes, which in all Heteronemertines are in direct continuity of tissue with the upper pair of principal lobes, cease to have this intimate connexion in the Metanemertini; and, although still constituted of (1) a ciliated duct, opening out externally, (2) nervous tissue surrounding it, and (3) histological elements distinctly different from the nervous, and most probably directly derived from the esophageal outgrowths, they are nevertheless here no longer constantly situated behind the upper brain-lobes and directly connected with them, but are found sometimes behind, sometimes beside and sometimes before the brain-lobes. Furthermore, they are here severed from the principal lobes and connected with them by one or more rather thick strings of nerve-fibres. In some cases, especially when the lobes lie before the brain, their distance from it, as well as the length of these nervous connexions, has considerably increased.
These curious neuro-glandular pits (fig. 1), absent in the Mesonemertine and one or two aberrant species, have been shown to possess large glandular cells at their base which secrete a mucus. The development of these organs, which in the Protonemertine are but grooves in the epidermis, not far removed from the similar cephalic slits of many Turbellaria, reaches its height in Drepanophorus. Here the pits split into two, one part ending in a sac lined with sensory epithelium, and embedded in nervous tissue, the other projecting backwards as a long, glandular, blind canal. The exit of these organs takes many shapes, of value in systematic work. Their function is still little understood. Two lateral, shallow pits occur on the side of the body about the level of the hinder end of the proboscis in some species of the genus Carinella, which are termed side-organs. These are capable of being everted, and are probably sensory in function (fig. 20, 17).
For the Heteronemertines arguments have been adduced to prove that here they have the physiological significance of a special respiratory apparatus for the central nervous tissue, which in all these forms is strongly charged with haemoglobin. The haemoglobin would by its pre-eminent properties of fixing oxygen, serve to furnish the nerve system, which more than any other requires a constant supply, with the necessary oxygen. Such could hardly be obtained in any other way by those worms that have no special respiratory apparatus, and that live in mud and under stones where the natural supply of freshly oxygenated sea-water is practically limited. Whether in the Metanemertines, where the blood fluid is often provided with haemoglobiniferous disks, the chief functions of the side organs may not rather be a sensory one needs further investigation.
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| Fig. 13. | Fig. 14. | |
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Figs. 13, 14.—Lateral views of head of a Heteronemertine (fig. 13) with longitudinal slit, and of a Metanemertine (fig. 14) with transverse groove and furrows. | ||
The exterior opening of the duct has been several times alluded to. In the Metanemertines it is generally situated towards the middle of a lateral transverse groove on either side of the head, as was noticed for Carinella, and as is also present in Polia. Generally a row of shorter grooves perpendicular to the first, and similarly provided with strong cilia, enlarges the surface of these furrows (fig. 14). In Valencinia there is nothing but a circular opening without furrow. In all Heteronemertines there is on each side of the head a longitudinal slit of varying length but generally considerable depth, in the bottom of which the dark red brain is very plainly visible by transparency. These slits are continued into the ciliated duct, being at the same time themselves very strongly ciliated. In life they are commonly rhythmically opened and shut by a wavy movement. They are the head slits (cephalic fissures, “Kopfspalten”) so characteristic of this subdivision (figs. 10 and 13).
With respect to the sense organs of the Nemertines, we find that eyes are of rather constant occurrence, although many Heteronemertines living in the mud appear to be blind. The more highly organized species have often very numerous eyes (Amphiporus, Drepanophorus), which are provided with a spherical refracting anterior portion, with a cellular “vitreous body,” with a layer of delicate radially arranged rods, with an outer sheath of dark pigment, and with a separate nerve-twig each, springing from a common or double pair of branches which leave the brain as n. optici, for the innervation of the eyes. Besides these more highly differentiated organs of vision, more primitive eyes are present in others down to simple stellate pigment specks without any refracting apparatus.
Organs of hearing in the form of capsules containing otoliths have only been very rarely observed, apparently only in Metanemertini.
As to the organ of touch, the great sensitiveness of the body has already been noticed, as well as the probable primary significance of the proboscis. Small tufts of tactile hairs or papillae are sometimes observed in small number at the tip of the head; sometimes longer hairs, apparently rather stiff, are seen on the surface, very sparingly distributed between the cilia, and hitherto only in a very limited number of small specimens. They may perhaps be considered as sensory.
Digestive System.—The anterior opening, the mouth, is situated ventrally, close to the tip of the head and in front of the brain in the Metanemertini, somewhat more backward and behind the brain in the other Nemertines. In most Heteronemertines it is found to be an elongated slit with corrugated borders; in the Metanemertines it is smaller and rounded; in Malacobdella and Akrostomum it, moreover, serves for the extrusion of the proboscis, which emerges by a separate dorsal opening just inside the mouth. The oesophagus is the anterior portion of the digestive canal; its walls are folded longitudinally, comparatively thick and provided with longitudinal muscular fibres. Two layers are specially obvious in its walls—the inner layer bordering the lumen being composed of smaller ciliated cells, the outer thicker one containing numerous granular cells and having a more glandular character. Outside the wall of the oesophagus a vascular space has been detected which is in direct continuity with the longitudinal blood-vessels. In certain cases, however, the walls of the oesophagus appear to be very closely applied to the muscular body-wall and this vascular space thereby considerably reduced.
The posterior portion of the intestine is specially characterized by the appearance of the intestinal diverticula horizontally and symmetrically placed right and left and opposite to each other.
In the Metanemertini there is a curious diverticulum of the intestine which stretches forward in the median line, ventral to the so-called stomach. It is at times sacculated, but its chief interest is that, as Lebedinsky[2] has shown, the tip of the caecum in embryonic life opens to the exterior as the blastopore. This subsequently closes up, and the newly-formed oesophagus and stomach open in the intestine above and behind it. It is a curious feature in Nemertines that the alimentary canal seldom contains traces of food and yet most of these worms are voracious. The food must be digested, absorbed and excreted with great rapidity. There is some evidence that in this group the ectoderm of the oesophagus is chiefly concerned with digestion, whereas the endoderm of the intestine is limited to the absorption of the soluble products.
Cases of asymmetry or irregularity in the arrangement of the
intestinal caeca, though sometimes occurring, are not normal. At
the tip of the tail, where the growth of the animal takes place, the caeca are always eminently regular. So they are throughout the
whole body in most of the Metanemertines. In Carinella they are
generally deficient and the intestine straight; in young specimens
of this species, however, they occur, though less regular and more in
the form of incipient foldings by which the digestive surface is,
increased. The inner surface of the intestinal caeca is ciliated, the
caeca themselves are
sometimes—especially in the
hindermost portion of the
body—of a considerably
smaller lumen than the
intermediate genital spaces;
sometimes, however, the
reverse is the case and in
both cases it is the smaller
lumen that appears enclosed
between and suspended by
the transverse fibres
constituting the muscular
dissepiments above mentioned.
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| Fig. 15 | |
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| Fig. 16 | |
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| Fig. 17 | |
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Figs. 15–17.—Diagrammatic sections to show disposition of internal organs in Carinella (Protonemertini), fig. 15, Heteronemertini, fig. 16, and Metanemertini, fig. 17. | |
| C, | Cellular portion of integument. |
| B, | Basement membrane. |
| A, | Circular muscular layer. |
| A′, | Longitudinal muscular layer. |
| A″, | Second circular (in Carinella). |
| A‴, | Second longitudinal (in Heteronemertini). |
| N, | Nervous layer. |
| LN, | Lateral nerves. |
| PS, | Cavity of proboscidian sheath (the sheath itself of varying thickness).
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| P, | Proboscis. |
| I, | Intestine. |
| LBv, Lateral blood-vessel. | |
| DBv, Dorsal blood-vessel. | |
| CT, | Connective tissue. |
The anus is situated terminally, the muscular body-wall through which the intestine must find its way outwards probably acting in this region the part of a sphincter. The lateral nerve stems mostly terminate on both sides in closest proximity to the anus; in certain species, however, they interfuse by a transverse connexion above the anus. The longitudinal blood-vessels do the same.
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Fig. 18.—Diagram of the circulatory apparatus in the anterior body-region of a Metanemertine. |
Circulatory apparatus.—The chief vessels are three longitudinal trunks, a median and two lateral ones. They are in direct connexion with each other both at the posterior and at the anterior end of the body. At the posterior end they communicate together by a T-shaped connexion in a simple and uniform way. Anteriorly there is a certain amount of difference in the arrangement. Whereas in the Metanemertines an arrangement prevails as represented in fig. 18, in the Heteronemertines the lateral stems, while entirely uniform all through the posterior portion of the body, no longer individually exist in the oesophageal region, but here dissolve themselves into a network of vascular spaces surrounding this portion of the digestive tract. The median dorsal vessel, however, remains distinct, but instead of continuing its course beneath the proboscidian sheath it is first enclosed by the ventral musculature of this organ, and still farther forwards it even bulges out longitudinally into the cavity of the sheath. Anteriorly it finally communicates with the lacunae just mentioned, which surround the oesophagus, bathe the posterior lobes of the brain, pass through the nerve ring together with the proboscidian sheath, and are generally continued in front of the brain as a lacunar space in the muscular tissue, one on each side.
Special mention must be made of the delicate transverse vessels regularly connecting the longitudinal and the lateral ones. They are metamerically placed, and belong to the same metamere as the digestive caeca, thus alternating with the generative sacs. The blood fluid does not flow in any definite direction; its movements are largely influenced by those of the muscular body-wall. It is colourless and contains definite corpuscles, which are round or elliptical, and in many Metanemertines are coloured red by haemoglobin, being colourless in other species. The circulatory system of Carinella is considerably different, being more lacunar and less restricted to definite vascular channels. Two lateral longitudinal lacunae form, so to say, the forerunners of the lateral vessels. A median longitudinal vessel and transverse connecting trunks have not as yet been detected. There are large lacunae in the head in front of the ganglia.
The vascular system is entirely closed. It contains a colourless fluid, with flat, oval nucleated corpuscles, as a rule colourless, but in some cases tinged with yellow or red haemoglobin. Its presence is one of the most distinctive features which separate the Nemertines from the Platyhelminthes. In origin the vascular system is due to a fusion of spaces which arise in the mesoblast of the larva. The blood is probably circulated by the general contraction of the whole animal, since it is very doubtful if there are any intrinsic muscles in the vessel-walls. Its function is less that of respiration than of conveying the digested food-products all over the body, and the excretory products to the nephridia, and doubtless it serves at times to assist in the extension and retraction of parts of the body. The vessels in the more highly-developed genera seem to be partly lacunae and partly true vessels with definite walls.
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Fig. 19.—Part of the excretory system lying on the lateral vessel of Drepanophorus spectabilis. (Magnified about 750.) 1, The longitudinal excretory canal; 2, one of the tags containing the flame-cells. |
Nephridia.—Associated with the lateral blood-vessels are the single pair of nephridia. Each consists of a more or less coiled, ciliated, longitudinal canal, which on its external surface gives origin to one or more transverse canals, which pass to the exterior and open a little way behind the mouth on the sides of the body. On its inner surface the longitudinal canal is adpressed to the lateral blood-vessel, and gives off a number of small, blind caeca or tags, each of which ends in a small clump of cells. These tags indent the blood-vessel. From their inner ends, projecting into the lumen of the tag, hangs a bunch of cilia, which forms the flickering “flame” so well known in the excretory apparatus of the Platyhelminthes and larval Annelids (fig. 19). There is no communication between the nephridia on one side and the other, but in Eupolia there are ducts opening into the alimentary canal as well as to the exterior, a condition of things which recalls what obtains in certain Oligochaetes. As a rule these organs only extend a short way along the anterior end of the body, a concentration which we may associate with the development of a vascular system to bring the products of excretion to a fixed spot. In Stichostemma, however, Montgomery[3] has described a series of nephridia lying all along the body, and each with a varying number of external pores. The excretory system is epiblastic in its origin.
The two external openings of the nephridia are situated sometimes more towards the ventral, at other times more towards the dorsal side. Even in the larger Heteronemertines these pores are only a few millimetres behind the mouth region. In transverse sections the nephridia can be shown to be generally situated in the region limited by (1) the proboscidian sheath, (2) the upper wall of the intestine, (3) the muscular body-wall. No trace of nephridia is found posterior to the oesophagus.
Generative System.—In the Nemertines the sexes are separate, with only very few exceptions (Tetrastemma hermaphroditica, Marion). The reproductive system is of the simplest, strongly contrasting with the complicated arrangements in the Platyhelminthes. A series of sacs lined with an epithelium, the proliferation of which gives rise to the ova or spermatozoa, alternate between the caeca of the intestine. When mature, each sac pushes out a process to the exterior, and this forms the genital duct. The line of the genital openings is usually dorsal to the lateral nerve. The whole sac, with its epithelial wall and its contained genital cells, arises ultimately from some of the parenchymatous cells of the body. The walls and contents in some forms arise simultaneously; in others the walls are first formed and their lining then proliferates. It has been pointed out that the cavity of the sacs corresponds in many particulars with the coelom of higher animals, and in Lebidinsky’s observations on the development there is some support to the view that a coelom exists. Montgomery has also described certain spaces which may be coelomic lying between the alimentary canal and the inner longitudinal layer of muscles in the Heteronemertini. The ova and spermatozoa, when mature, present no peculiarities. As the ova are in many species deposited in a gelatinous tube secreted by the body-walls, in which they are arranged (three or more together) in flask-shaped cavities, impregnation must probably take place either before or at the very moment of their being deposited. The exact mode has not yet been noticed.
Prosorhocmus claparedii is a viviparous form.
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Fig. 20.—Anterior end of a Carinella, partly diagrammatic. Magnified. (From Bürger.) 1, Opening of proboscis; 2, cephalic glands running to frontal organ; 3, dorsal commissure of brain; 4, cerebral organ; 5, upper dorsal nerve; 6, under dorsal nerve; 7, rhynchocoelic blood-vessel; 8, fore-gut; 9, rhynchocoel; 10, nerve to proboscis; 11, proboscis; 12, genital sac; 13, genital pore; 14, mid-gut; 15, circular nerves; 16, pore of excretory system; 17, lateral organ; 18, excretory canal; 19, lateral vessel; 20, lateral nerve; 21, oesophageal nerve; 22, mouth; 23, ventral ganglion of brain; 24, dorsal ganglion of brain; 25, rhynchodaeum. |
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| Fig. 21.—Pilidium-larva. | |
| B, | Bunch of cilia or flagellum.
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| oe, | Oesophagus. |
| st, | Stomach. |
| cs, | Oesophageal outgrowth for lateral organ.
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| am, | Amnion. |
| pr.d., Prostomial disk. | |
| po.d., Metastomial disk. | |
Development.—The embryology of the Nemertines offers some very remarkable peculiarities. Our knowledge of the development of the most primitive forms is scanty. Both Hetero- and Metanemertini have been more exhaustively studied than the other two groups, the first, as was noticed above, being characterized by peculiar larval forms, the second developing without metamorphosis.
The larva of Cerebratulus is called the pilidium. In exterior shape it resembles a helmet with spike and ear-lobes, the spike being a strong and long flagellum or a tuft of long cilia, the ear-lobes lateral ciliated appendages (fig. 21). It encloses the primitive alimentary tract. Two pairs of invaginations of the skin, which originally are called the prostomial and metastomial disks, grow round the intestine, finally fuse together, and form the skin and muscular body-wall of the future Nemertine, which afterwards becomes ciliated, frees itself from the pilidium investment and develops into the adult worm without further metamorphosis.
The eggs of these species are not enveloped by such massive gelatinous strings as are those of the genus Lineus. In the latter we find the young Nemertines crawling about after a period of from six to eight weeks, and probably feeding upon a portion of this gelatinous substance, which is found to diminish in bulk. In accordance with these more sedentary habits during the first phases of life, the characteristic pilidium larva, which is so eminently adapted for a pelagic existence, appear to have been reduced to a close-fitting exterior layer of cells, which is stripped off after the definite body-wall of the Nemertine has similarly originated out of four ingrowths from the primary epiblast. To this reduced and sedentary pilidium the name of “larva of Desor” has been given.
In the Metanemertini, as far as they have been investigated, a direct development without metamorphosis has been observed. It appears probable that this is only a further simplification of the more complicated metamorphosis described above.
As to the development of the different organs, there is still much that remains doubtful. The hypoblast in some forms originates by invagination, in others by delamination. The proboscis is an invagination from the epiblast; the proboscidian sheath appears in the mesoblast, but is perhaps originally derived from the hypoblast. The origin of the lateral organs has already been noticed; that of the nerve system is essentially epiblastic.
Affinities.—The position of the Nemertines in the animal kingdom is now looked upon as more isolated than was formerly thought, and recent writers have been inclined to treat them as a separate hylum. Whether this view be adopted or not, and whether the Turbellaria be regarded as nearly related or only remotely connected, there can be little doubt that the Nemertines resemble the Turbellaria more nearly than they do any other group of animals. Bürger even goes so far as to homologize the proboscis with the Turbellarian pharynx, and he sums up their relationship to the Annelids by the statement that to a certain extent the Nemertines represent Turbellaria which in the course of time have copied certain features of an Annelid character.
Literature.—J. Barrois, “Recherches sur l'embryologie des Némertes,” Annales des Sc. Naturelles, vi. (1877); O. Bütschli, “Einige Bemerkungen zur Metamorphose des Pilidium,” Archiv für Naturgeschichte (1873); L. von Graff, Monographie der Turbellarien (1882); A. A. W. Hubrecht, “Untersuchungen über Nemertinen a. d. Golf von Neapel,” Niederl. Archiv für Zoologie, ii.; Id., “The Genera of European Nemerteans critically revised,” Notes from the Leyden Museum (1879); Id., “Zur Anatomie u. Physiologie d. Nervensystems d. Nemertinen,” Verh. kon. Akad. v. Wetensch. (Amsterdam, 1880), vol. xx.; Id., “The Peripheral Nervous System of the Palaeo- and Schizonemertini, one of the layers of the Body-wall,” Quart. Journal of Micr. Science, vol. xx.; Id., “On the Ancestral Forms of the Chordata,” Ib. (July 1883); W. Keferstein, “Untersuchungen über niedere Seethiere,” Zeitschr. f. wissensch. Zool. vol. xii. (1863); J. von Kennel, “Beiträge zur Kenntniss der Nemertinen,” Arbeiten a. d. zool.-zoot. Instit. ii. (Würzburg, 1878); W. C. MacIntosh, A Monograph of British Annelida: I. Nemerteans (Ray Society, 1873-1874); A. F. Marion, “Recherches sur les animaux inférieurs du Golfe de Marseille,” Ann. des Sc. Nat. (1873); E. Metschnikoff, “Studien über die Entwickelung der Echinodermen und Nemertinen,” Mém. de l'Acad. Imp. de St Petersb. xiv. (1869); Max Schultze, Beiträge zur Naturgeschichte der Turbellarien (Greifswald, 1851) and Zeitschr. für wissensch. Zool. iv. (1852), p. 178; W. B. Benham, Quart. Journ. Micr. Sci. xxxix. (1896), p. 19; A. Brown, Proc. Roy. Soc. lxi. (1897), p. 28; O. Bürger, Zeit. f. wiss. Zool. l. (1890), p. 1; Id., Mitt. Zool. St Neapel, x. (1891), p. 206; Id., Zeit. f. wiss. Zool. liii. (1892), p. 322; Id., Verh. Deutsch. zool. Gesellsch. (1893); Id., Fauna u. Flora d. Golfe d. Neapel, Monograph 22 (1895); A. Dendy, Proc. Roy. Soc. Victoria (n.s.), iv. (1892), p. 85, v. p. 127 (1891-1892); B. Haller, Arb. Zool. Inst. Wien, viii. (1889), p. 276; A. A. W. Hubrecht, “Challenger” Reports, xix. (1887); L. Joubin, Arch. Zool. Exper. (2), viii. (1890), p. 461; Id., “Nemertines,” in Blanchard’s Traité de zoologie (1894); J. N. Lebedinsky, Arch. Mikr. Anat. xlix. (1897), p. 503: T. H. Montgomery, Zool. Anzeig. xvii. (1894), pp. 298, 301; Id., Zeit. f. wiss. Zool. lix. (1895), p. 83; Id., Zool. Jahrb. (Anat.) x. (1897), p. 1; A. E. Verrill, Trans. Connecticut Acad. New Haven, viii. (1892), p. 382; D. Bergendal, Zool. Anzeiger, xxiii. (1900), p. 313; W. R. Coe, Zool. Jahrb. (Anat.) xii. (1899), p. 425; Id., Trans. Connect. Acad. ix. (1895), p. 479; Id., Proc. Wash. Acad. iii. (1901), p. 1; T. H. Montgomery, Journ. Morph., xiii. (1897) p. 381; Id., Zool. Jahrb. (Anat.) x. (1897), p. 265; R. C. Punnett, Quart. Journ. Mic. Sc. xliv. (1900), p. 111; Id., Willey’s Zool. Results, pt. v. (1900), p. 569; Id., Quart. Journ. Mic. Sc. xliv. (1901), p. 547; Staub, Semon’s Forschungsreisen (5 Bd., 1900); C. B. Thompson, Zool. Anzeiger, xxiii. (1900), pp. 151, 627; C. B. Wilson, Quart. Journ. Mic. Sc. xliii. (1900), p. 97. (A. A. W. H.; A. E. S.)