Encyclopaedia Britannica, 11th Edition, "Electrostatics" to "Engis" / Volume 9, Slice 3

by Various

In the Echinodermata, Enteropneusta and one or two other groups, it is not possible, in the present state of knowledge, to bring the mouth into relation with the blastopore, nor can the blastopore be shown to be a perforation of the neural surface. For the Echinoderms, at any rate, this fact loses some of the importance which might at first sight be attributed to it when the remarkable organization of the adult and the sharp contrast which exists between it and the larva is remembered. In some Annelids the central nervous system remains throughout life as part of the outer epidermis, but as a general rule it becomes separated from the epidermis and embedded in the mesodermal tissues. The mode in which this separation is effected varies according to the form and structure of the central nervous system. In the Vertebrata, in which this organ has the form of a tube extending along the dorsal surface of the body, it arises as a groove of the medullary plate, which becomes constricted into a canal. The wall of this canal consists of ectoderm, which at an earlier stage formed part of the outer surface of the body, but which after invagination thickens, to give rise to the epithelial lining of the canal and to the nervous tissue which forms the bulk of the canal wall. The fact that the blastopore remains open at the hind end of the medullary plate explains to a certain extent the peculiar relation which always exists in the embryo between the hind end of the neural and alimentary canals. This communication between the hind end of the neural tube and the gut is one of the most remarkable and constant features of the Vertebrate embryo. As has been pointed out, it is not altogether unintelligible when we remember the relation of the blastopore to the medullary plate of the earlier stage, but to give a complete explanation of it is, and probably always will be, impossible. It is no doubt the impress of some remarkable larval condition of the blastopore of a stage of evolution now long past.

In Ceratodus the open part of the blastopore is enclosed by the medullary folds, as in Lepidosiren, and probably persists as the anus, the portion of the folds around the anus undergoing atrophy (Semon, Zool. Forschungsreisen in Australien, 1893, Bd. i. p. 39). In Urodeles the blastopore persists as anus, so far as is known, but the relation to the medullary folds has not been noticed. The same may be said of Petromyzon (A.E. Shipley, Quart. Journ. Mic. Sci. xxviii., 1887).

The nerve tube of the Vertebrata at a certain early stage of the embryo becomes bent ventralwards in its anterior portion, in such a manner that the anterior end, which is represented in the adult by the infundibulum, comes to project Cranial flexure. backwards beneath the mid-brain. This bend, which is called the cranial flexure, takes place through the mid-brain, so that the hind-brain is unaffected by it. The cranial flexure is not, however, confined to the brain: the anterior end of the notochord, which at first extends almost to the front end of the nerve tube (this extension, which is quite obvious in the young embryo of Elasmobranchs, becomes masked in the later stages by the extraordinary modifications which the parts undergo), is also affected by it. Moreover, it affects even other parts, as may be seen by the oblique, almost antero-posterior, direction of the anterior gill slits as compared with the transverse direction of those behind. No satisfactory explanation has ever been offered of the cranial flexure. It is found in all Vertebrates, and is effected at an early stage of the development. In the later stages and in the adult it ceases to be noticeable, on account of an alteration of the relative sizes of parts of the brain. This is due almost entirely to the enormous growth of the cerebral vesicle, which is an outgrowth of the dorsal wall of the fore-brain just short of its anterior end. The anterior end of the fore-brain remains relatively small throughout life as the infundibulum, and the junction of this part of the fore-brain with the part which is so largely developed, as the rudiment of the cerebrum, is marked by the attachment of the optic chiasma. The optic nerve, indeed, is morphologically the first cranial nerve, the olfactory being the second; both are attached to what is morphologically the dorsal side of the nerve tube. The morphological anterior end of the central nerve tube is the point of the infundibulum which is in contact with the pituitary body. While on the subject of the cranial flexure, it may be pointed out that there is a similar downward curve of the hind end of the nervous axis, which leads into the hind end of the enteron. If it be supposed that originally there was a communication between the infundibulum and pituitary body, then the ventral flexure found at both ends of the nerve axis would originally have had the same result, namely, of placing the neural and alimentary canals in communication. Moreover, the mouth would have had much the same relation to this imaginary anterior neurenteric canal that the anus has to the actual posterior one.

In Amphioxus and the Tunicata the early development of the central nervous system is very much like that of the Vertebrata, but the later stages are simpler, being without the cranial flexure. The Tunicata are remarkable for the fact that the nervous system, though at first hollow, becomes quite solid in the adult. In Balanoglossus the central nervous system is in part tubular, the canal being open at each end. It arises, however, by delamination from the ectoderm, the tube being a secondary acquisition. This is probably due to a shortening of development, for the same feature is found in some Vertebrata (Teleostei, Lepidosteus, &c.), where the central canal is secondarily hollowed out in the solid keel-like mass which is separated from the ectoderm. Parts of the central nervous system arise by invagination in other groups; for instance, the cerebral ganglia of Dentalium are formed from the walls of two invaginations of ectoderm, which eventually disappear at the anterior end of the body (A. Kowalevsky, Ann. Mus. Hist. Nat. Marseilles, “Zoology,” vol. i.). In Peripatus the cerebral ganglia arise in a similar way, but in this case the cavities of the invagination become separated from the skin and persist as two hollow appendages on the lower side of the cerebral ganglia. In other Arthropods the cerebral ganglia arise in a similar way, but the invaginations disappear in the adult. In Nemertines the cerebral ganglia contain a cavity which communicates with the exterior by a narrow canal. Finally, in certain Echinodermata the ventral part of the central nervous system arises by the invagination of a linear streak of ectoderm, the cavity of the invagination persisting as the epineural canal.

Although the central nervous system is almost always developed from the ectoderm of the embryo, the same cannot be said of the peripheral nerve trunks. These structures arise from the mesoblastic reticulum already described Peripheral nervous system. (Sedgwick, Quart. Journ. Mic. Sci. xxxvii. 92). Inasmuch as this reticulum is perfectly continuous with the precisely similar though denser tissue in the ectoderm and endoderm, it may well be that a portion of the nerve trunks should be described as being ectodermal and endodermal in origin, though the bulk of them are undoubtedly formed from that portion of the reticulum commonly described as mesoblastic. But, however that may be, the tissue from which the great nerve trunks are developed is continuous on all sides with a similar tissue which pervades all the organs of the body, and in which the nuclei of these organs are contained.

In the early stages of development this tissue is very sparse and not easily seen. It would appear, indeed, that it is of a very delicate texture and readily destroyed by reagents. It is for this reason that the layers of the Vertebrate embryo are commonly represented as being quite isolated from one another, and that the medullary canal is nearly always represented as being completely isolated at certain stages from the surrounding tissues. In reality the layers are all connected together by this delicate tissue—in a sparse form, it is true—which not only extends between them, but also in a denser and more distinct form pervades them. In the germinal layers themselves, and in the organs developing from them, this tissue is in the young stages almost entirely obscured by the densely packed nuclei which it contains. For instance, in the wall of the medullary canal in the Vertebrate embryo, in the splanchnic and somatic layers of mesoderm of the same embryo, and in the developing nerve cords of the Peripatus embryo, the nuclei are at first so densely crowded together that it is almost impossible to see the protoplasmic framework in which they rest, but as development proceeds this extra-nuclear tissue becomes more largely developed, and the nuclei are forced apart, so that it becomes visible and receives various names according to its position. In the wall of the medullary canal of the Vertebrate embryo, on the outside of which it becomes especially conspicuous in certain places, and on the dorsal side of the developing nerve cords of the Peripatus embryo, it constitutes the white matter of the developing nerve cord; in the mesoblastic tissue outside, where it at the same time becomes more conspicuous (Sedgwick, “Monograph of the Development of Peripatus capensis,” Studies from the Morph. Lab. of the University of Cambridge, iv., 1889, p. 131), it forms the looser network of the mesoblastic reticulum; and connecting the two, in place of the few and delicate strands of this tissue of the former stage, there are at certain places well-marked cords of a relatively dense texture, with the meshes of the reticulum elongated in the direction of the cord. This latter structure is an incipient nerve trunk. It can be traced outwards into the mesoblastic reticulum, from the strands of which it is indeed developed, and with which it is continuous not only at its free end, but also along its whole course. In this way the nerve trunks are developed—by a gathering up, so to speak, of the fibres of the reticulum into bundles. These bundles are generally marked by the possession of nuclei, especially in their cortical parts, which become no doubt the nuclei of the nerve sheath, and, in the neighbourhood of the ganglia, of nerve cells. From this account of the early development of the nerves, it is apparent that they are in their origin continuous with all the other tissues of the body, with that of the central nervous system and with that which becomes transformed into muscular tissue and connective and epithelial tissues. All these tissues are developed from the general reticulum, which in the young embryo can be seen to pervade the whole body, not being confined to the mesoderm, but extending between the nuclei of the ectoderm and endoderm, and forming the extra-nuclear, so-called cellular, protoplasm of those layers. Moreover, it must be remarked that in the stages of the embryo with which we are here concerned the so-called cellular constitution of the tissues, which is such a marked feature of the older embryo and adult, has not been arrived at. It is true, indications of it may be seen in some of the earlier-formed epithelia, but of nerve cells, muscular cells, and many kinds of gland cells no distinct signs are yet visible. This remark particularly applies to nerve cells, which do not make their appearance until a much later stage—not, indeed, until some time after the principal nerve trunks and ganglia are indicated as tracts of pale fibrous substance and aggregations of nuclei respectively.

The embryos of Elasmobranchs—particularly of Scyllium—are the best objects in which to study the development of nerves. In many embryos it is difficult to make out what happens, because the various parts of the body remain so close together that the process is obscured, and the loosening of the mesoblastic nuclei is deferred until after the nerves have begun to be differentiated. The process may also be traced in the embryos of Peripatus, where the main features are essentially similar to those above described (op. cit. p. 131). The development of the motor nerves has been worked out in Lepidosiren by J. Graham Kerr (Trans. Roy. Soc. of Edinburgh, 41, 1904. p. 119).

To sum up, the development of nerves is not, as has been recently urged, an outgrowth of cell processes from certain cells, but is a differentiation of a substance which was already in position, and from which all other organs of the body have been and are developed. It frequently happens that the young nerve tracts can be seen sooner near the central organ than elsewhere, but it is doubtful if any importance can be attached to this fact, since it is not constantly observed. For instance, in the case of the third nerve of Scyllium the differentiation appears to take place earliest near the ciliary ganglion, and to proceed from that point to the base of the mid-brain.

There are two main methods in which new organs are developed. In the one, which indicates the possibility of physiological continuity, the organ arises by the direct modification of a portion of a pre-existing organ; Coelom. the development of the central nervous system of the Vertebrata from a groove in the embryonic ectoderm may be taken as an example of this method. In the other method there is no continuity which can be in any way interpreted as physiological; a centre of growth appears in one of the parts of the embryo, and gives rise to a mass of tissue which gradually shapes itself into the required organ. The development of the central nervous system in Teleosteans and in other similar exceptional cases may be mentioned as an example of the second plan. Such a centre of growth is frequently called a blastema, and consists of a mass of closely packed nuclei which have arisen by the growth-activity of the nuclei in the neighbourhood. The coelom, an organ which is found in the so-called coelomate animals, and which in the adult is usually divided up more or less completely into three parts, namely, body-cavity, renal organs, generative glands, presents in different animals both these methods of development. In certain animals it develops by the direct modification of a part of the primitive enteron, while in others it arises by the gradual shaping of a mass of tissue which consists of a compact mass of nuclei derived by nuclear proliferation from one or more of the pre-existing tissues of the body. Inasmuch as the first rudiment of the coelom nearly always makes its appearance at an early stage, when the ectoderm and endoderm are almost the only tissues present, and as it then bulks relatively very large and frequently contains within itself the potential centres of growth of other organs, e.g. mesenchymal organs (see above), it has come to be regarded by embryologists as being the forerunner of all the so-called mesodermal organs of the body, and has been dignified with the somewhat mysterious rank which attaches to the conception of a germinal layer. Its prominence and importance at an early stage led embryologists, as has already been explained, to overlook the fact that although some of the centres of growth for the formation of other non-coelomic mesodermal organs and tissues may be contained within it, all are not so contained, and that there are centres of mesodermal growth still left in the ectoderm and endoderm after its establishment. If these considerations, and others like them, are correct, it would seem to follow that the conception implied by the word mesoderm has no objective existence, that the tissue of the embryo called mesoderm, though sometimes mainly the rudiment of the coelom, is often much more than this, and contains within itself the rudiment of many, sometimes of all, of the organs appertaining to the mesenchyme. In thus containing within itself the potential centres of growth of other organs and tissues which are commonly ranked as mesodermal, it is not different from the rudiments of the two other organs already formed, namely, the ectoderm and endoderm; for these contain within themselves centres of growth for the production of so-called mesodermal tissues, as witness the nerve-crest of Vertebrata, the growing-point of the pronephric duct, and the formation of blood-vessels from the hypoblast described for some members of the same group.

In Echinodermata, Amphioxus, Enteropneusta, and a few other groups, the coelom develops from a portion or portions of the primitive enteron, which eventually becomes separated from the rest and forms a variable number of closed sacs lying between the gut and the ectoderm. The number of these sacs varies in different animals, but the evidence at present available seems to show that the maximum number is five—an unpaired one in front and two pairs behind—and, further, that if a less number of sacs is actually separated from the enteron, the rule is for these sacs so to divide up that they give rise to five sacs arranged in the manner indicated. The Enteropneusta present us with the clearest case of the separation of five sacs from the primitive enteron (W. Bateson, Quart. Journ. Mic. Sci. xxiv., 1884). In Amphioxus, according to the important researches of E.W. MacBride (Quart. Journ. Mic. Sci. xl. 589), it appears that a similar process occurs, though it is complicated by the fact that the sacs of the posterior pair become divided up at an early stage into many pairs. In Phoronis there are indications of the same phenomenon (A.T. Masterman, Quart. Journ. Mic. Sci. xliii. 375). In the Chaetognatha a single sac only is separated from the enteron, but soon becomes divided up. In the Brachiopoda one pair of sacs is separated from the enteron, but our knowledge of their later history is not sufficient to enable us to say whether they divide up into the typically arranged five sacs. In Echinodermata the number of sacs separated from the enteron varies from one to three; but though the history of these shows considerable differences, there are reasons to believe that the typical final arrangement is one unpaired and two paired sacs. But however many sacs may arise from the primitive enteron, and however these sacs may ultimately divide up and arrange themselves, the important point of development common to all these animals, about which there can be no dispute, is that the coelom is a direct differentiation of a portion of the enteron.