The wide divarication of the ventral nerve cords in the embryo renders it easy to prove that there is no median invagination of epiblast between them, and supports Kleinenberg's observations on Lumbricus as to the absence of this invagination. I have further satisfied myself as to the absence of such an invagination in Peripatus. It is probable that Hatschek and other observers who have followed him are mistaken in affirming the existence of such an invagination in either the Chætopoda or the Arthropoda.

The observations recorded in this paper on the yolk cells and their derivations are, on the whole, in close harmony with the observations of Dohrn, Bobretzky, and Graber, on Insects. They shew, however, that the first formed mesoblastic plate does not give rise to the whole of the mesoblast, but that during the whole of embryonic life the mesoblast continues to receive accessions of cells derived from the cells of the yolk.

Araneina.

1. Balbiani, “Mémoire sur le Développement des Araneides,” Ann. Sci. Nat., series v, Vol. XVII. 1873.

2. J. Barrois, “Recherches s. l. Développement des Araignées,” Journal de l'Anat. et de la Physiol., 1878.

3. E. Claparède, Recherches s. l'Evolution des Araignées, Utrecht, 1860.

4. Herold, De Generatione Araniorum in Ovo, Marburg, 1824.

5. H. Ludwig, “Ueb. d. Bildung des Blastoderm bei d. Spinnen,” Zeit. f. wiss. Zool., Vol. XXVI. 1876.

Plate 30.

Complete List of Reference Letters.

ch. Cheliceræ. ch.g. Ganglion of cheliceræ. c.l. Caudal lobe. p.c. Primitive cumulus. pd. Pedipalpi. pr.l. Præoral lobe. pp¹. pp². etc. Provisional appendages. sp. Spinnerets. st. Stomodæum.

I-IV. Ambulatory appendages. 1-16. Postoral segments.

Fig. 1. Ovum, with primitive cumulus and streak proceeding from it.

Fig. 2. Somewhat later stage, in which the primitive cumulus is still visible. Near the opposite end of the blastoderm is a white area, which is probably the rudiment of the procephalic lobe.

Fig. 3a and 3b. View of an embryo from the ventral surface and from the side when six segments have become established.

Fig. 4. View of an embryo, ideally unrolled, when the first rudiments of the appendages become visible.

Fig. 5. Embryo ideally unrolled at the stage when all the appendages have become established.

Fig. 6. Somewhat older stage, when the limbs begin to be jointed. Viewed from the side.

Fig. 7. Later stage, viewed from the side.

Fig. 7a. Same embryo as fig. 7, ideally unrolled.

Figs. 8a and 8b. View from the ventral surface and from the side of an embryo, after the ventral flexure has considerably advanced.

Fig. 9. Somewhat older embryo, viewed from the ventral surface.

PLATES 31 AND 32.

Complete List of Reference Letters.

Ig-IVg. Ganglia of ambulatory limbs. 1-16. Postoral segments.

Fig. 10. Section through an ovum, slightly younger than fig. 1. Shewing the primitive cumulus and the columnar character of the cells of one half of the blastoderm.

Fig. 11. Section through an embryo of the same age as fig. 2. Shewing the median thickening of the blastoderm.

Fig. 12. Transverse section through the ventral plate of a somewhat older embryo. Shewing the division of the ventral plate into epiblast and mesoblast.

Fig. 13. Section through the ventral plate of an embryo of the same age as fig. 3, shewing the division of the mesoblast of the ventral plate into two mesoblastic bands.

Fig. 14. Transverse section through an embryo of the same age as fig. 5, passing through an abdominal segment above and a thoracic segment below.

Fig. 15. Longitudinal section slightly to one side of the middle line through an embryo of the same age.

Fig. 16. Transverse section through the ventral plate in the thoracic region of an embryo of the same age as fig. 7.

Fig. 17. Transverse section through the procephalic lobes of an embryo of the same age. gr. Section of hemicircular groove in procephalic lobe.

Fig. 18. Transverse section through the thoracic region of an embryo of the same age as fig. 8.

Fig. 19. Section through the procephalic lobes of an embryo of the same age.

Fig. 20a, b, c, d, e. Five sections through an embryo of the same age as fig. 9. a and b are sections through the procephalic lobes, c through the front part of the thorax. d cuts transversely the posterior parts of the thorax, and longitudinally and horizontally the ventral surface of the abdomen. e cuts the posterior part of the abdomen longitudinally and horizontally, and shews the commencement of the mesenteron.

Fig. 21. Longitudinal and vertical section of an embryo of the same age. The section passes somewhat to one side of the middle line, and shews the structure of the nervous system.

Fig. 22. Transverse section through the dorsal part of the abdomen of an embryo of the same stage as fig. 9.

By treating with nitric acid and clarifying by oil of cloves, and subsequently removing one half of the body so as to expose the spinal cord in sitû, the origin and distribution of the posterior nerves is very clearly exhibited. But I have failed to detect any trace of the anterior nerve-roots. Horizontal section, which ought also to bring them clearly into view, failed to shew me anything which I could interpret as such. I agree with Schneider that a process of each muscle-plate is prolonged up to the anterior border of the spinal cord, but I can find no trace of a connection between it and the cord.

Schneider has represented a transverse section in which the anterior nerves are figured. I am very familiar with an appearance in section such as that represented in his figure, but I satisfied myself when I previously studied the nerves in Amphioxus, that the body supposed to be a nerve by Schneider was nothing else than part of the intermuscular septum, and after re-examining my sections I see no reason to alter my view.

XIX. Address to the Department of Anatomy and Physiology of ohe British Association, 1880.

In the spring of the present year, Professor Huxley delivered an address at the Royal Institution, to which he gave the felicitous title of 'The coming of age of the origin of species.' It is, as he pointed out, twenty-one years since Mr Darwin's great work was published, and the present occasion is an appropriate one to review the effect which it has had on the progress of biological knowledge.

There is, I may venture to say, no department of biology the growth of which has not been profoundly influenced by the Darwinian theory. When Messrs Darwin and Wallace first enunciated their views to the scientific world, the facts they brought forward seemed to many naturalists insufficient to substantiate their far-reaching conclusions. Since that time an overwhelming mass of evidence has, however, been rapidly accumulating in their favour. Facts which at first appeared to be opposed to their theories have one by one been shewn to afford striking proofs of their truth. There are at the present time but few naturalists who do not accept in the main the Darwinian theory, and even some of those who reject many of Darwin's explanations still accept the fundamental position that all animals are descended from a common stock.

The law of variation is in a certain sense opposed to the law of heredity. It asserts that the resemblance which offspring bear to their parents is never exact. The contradiction between the two laws is only apparent. All variations and modifications in an organism are directly or indirectly due to its environments; that is to say, they are either produced by some direct influence acting upon the organism itself, or by some more subtle and mysterious action on its parents; and the law of heredity really asserts that the offspring and parent would resemble each other if their environments were the same. Since, however, this is never the case, the offspring always differ to some extent from the parents. Now, according to the law of heredity, every acquired variation tends to be inherited, so that, by a summation of small changes, the animals may come to differ from their parent stock to an indefinite extent.

The general nature of the methods of reasoning employed by embryologists, who accept the Darwinian theory, is exemplified by the instance just given. If this method is a legitimate one, and there is no reason to doubt it, we ought to find that animals, in the course of their development, pass through a series of stages, in each of which they resemble one of their remote ancestors; but it is to be remembered that, in accordance with the law of variation, there is a continual tendency to change, and that the longer this tendency acts the greater will be the total effect. Owing to this tendency, we should not expect to find a perfect resemblance between an animal, at different stages of its growth, and its ancestors; and the remoter the ancestors, the less close ought the resemblance to be. In spite, however, of this limitation, it may be laid down as one of the consequences of the law of inheritance that every animal ought, in the course of its individual development, to repeat with more or less fidelity the history of its ancestral evolution.

While, for the reasons just stated, it is not generally possible to prove by direct observation that existing forms in their embryonic state repeat the characters of their ancestors, there is another method by which the truth of this proposition can be approximately verified.

A comparison of recent and fossil forms shews that there are actually living at the present day representatives of a considerable proportion of the groups which have in previous times existed on the globe, and there are therefore forms allied to the ancestors of those living at the present day, though not actually the same species. If therefore it can be shewn that the embryos of existing forms pass through stages in which they have the characters of more primitive groups, a sufficient proof of our proposition will have been given.

That such is often the case is a well-known fact, and was even known before the publication of Darwin's works. Von Baer, the greatest embryologist of the century, who died at an advanced age but a few years ago, discussed the proposition at considerable length in a work published between the years 1830 and 1840. He came to the conclusion that the embryos of higher forms never actually resemble lower forms, but only the embryos of lower forms; and he further maintained that such resemblances did not hold at all, or only to a very small extent, beyond the limits of the larger groups. Thus he believed that, though the embryos of Vertebrates might agree amongst themselves, there was no resemblance between them and the embryos of any invertebrate group. We now know that these limitations of Von Baer do not hold good, but it is to be remembered that the meaning now attached by embryologists to such resemblances was quite unknown to him.

These preliminary remarks will, I trust, be sufficient to demonstrate how completely modern embryological reasoning is dependent on the two laws of inheritance and variation, which constitute the keystones of the Darwinian theory.

The discoveries recently made in organogeny, or the genesis of organs, have been quite as striking, and in many respects even more interesting, than those in phylogeny, and I propose devoting the remainder of my address to a history of results which have been arrived at with reference to the origin of the nervous system.

To render clear the nature of these results I must say a few words as to the structure of the animal body. The body is always built of certain pieces of protoplasm, which are technically known to biologists as cells. The simplest organisms are composed either of a single piece of this kind, or of several similar pieces loosely aggregated together. Each of these pieces or cells is capable of digesting and assimilating food, and of respiring; it can execute movements, and is sensitive to external stimuli, and can reproduce itself. All the functions of higher animals can, in fact, be carried on in this single cell. Such lowly organized forms are known to naturalists as the Protozoa. All other animals are also composed of cells, but these cells are no longer complete organisms in themselves. They exhibit a division of labour: some carrying on the work of digestion; some, which we call nerve-cells, receiving and conducting stimuli; some, which we call muscle-cells, altering their form—in fact, contracting in one direction—under the action of the stimuli brought to them by the nerve-cells. In most cases a number of cells with the same function are united together, and thus constitute a tissue. Thus the cells which carry on the work of digestion form a lining membrane to a tube or sack, and constitute a tissue known as a secretory epithelium. The whole of the animals with bodies composed of definite tissues of this kind are known as the Metazoa.

A considerable number of early developmental processes are common to the whole of the Metazoa.

There is every reason to think that Lankester and Haeckel were quite justified in concluding that a form more or less like that just described was the ancestor of the Metazoa; but the further speculations contained in their essays as to the origin of this form from the Protozoa can only be regarded as suggestive feelers, which, however, have been of great importance in stimulating and directing embryological research. It is, moreover, very doubtful whether there are to be found in the developmental histories of most animals any traces of this gastræa ancestor, other than the fact of their passing through a stage in which the cells are divided into two germinal layers.

The key to the nature of the two germinal layers is to be found in Huxley's comparison between them, and the two layers in the fresh-water polype and the sea-anemone. The epiblast is the primitive skin, and the hypoblast is the primitive epithelial wall of the alimentary tract.

In the whole of the polype group, or Cœlenterata, the body remains through life composed of the two layers, which Huxley recognized as homologous with the epiblast and hypoblast of the Vertebrata; but in all the higher Metazoa a third germinal layer, known as the mesoblast, early makes its appearance between the two primary layers. The mesoblast originates as a differentiation of one or of both the primary germinal layers; but although the different views which have been held as to its mode of origin form an important section of the history of recent embryological investigations, I must for the moment confine myself to saying that from this layer there take their origin—the whole of the muscular system, of the vascular system, and of that connective-tissue system which forms the internal skeleton, tendons, and other parts.

In any animal in which there is no distinct nervous system, it is obvious that the general surface of the body must be sensitive to the action of its surroundings, or to what are technically called stimuli. We know experimentally that this is so in the case of the Protozoa, and of some very simple Metazoa, such as the freshwater Polype or Hydra, where there is no distinct nervous system. The skin or epidermis of the ancestor of the Metazoa was no doubt similarly sensitive; and the fact of the nervous system being derived from the epiblast implies that the functions of the central nervous system, which were originally taken by the whole skin, became gradually concentrated in a special part of the skin which was step by step removed from the surface, and finally became a well-defined organ in the interior of the body.

What were the steps by which this remarkable process took place? How has it come about that there are nerves passing from the central nervous system to all parts of the skin, and also to the muscles? How have the arrangements for reflex actions arisen by which stimuli received on the surface of the body are carried to the central part of the nervous system, and are thence transmitted to the appropriate muscles, and cause them to contract? All these questions require to be answered before we can be said to possess a satisfactory knowledge of the origin of the nervous system. As yet, however, the knowledge of these points derived from embryology is imperfect, although there is every hope that further investigation will render it less so. Fortunately, however, a study of comparative anatomy, especially that of the Cœlenterata, fills up some of the gaps left from our study of embryology.

Another important fact shewn by embryology is that the central nervous system, and percipient portion of the organs of special sense, are often formed from the same part of the primitive epidermis. Thus, in ourselves and in other vertebrate animals the sensitive part of the eye, known as the retina, is formed from two lateral lobes of the front part of the primitive brain. The crystalline lens and cornea of the eye are, however, subsequently formed from the skin.

The same is true for the peculiar compound eyes of crabs or Crustacea. The most important part of the central nervous system of these animals is the supra-œsophageal ganglia, often known as the brain, and these are formed in the embryo from two thickened patches of the skin at the front end of the body. These thickened patches become gradually detached from the surface, remaining covered over by a layer of skin. They then constitute the supra-œsophageal ganglia; but they form not only the ganglia, but also the rhabdons or retinal elements of the eye—the parts in fact which correspond to the rods and cones in our own retina. The layer of epidermis or skin which lies immediately above the supra-œsophageal ganglia becomes gradually converted into the refractive media of the crustacean eye. A cuticle which lies on its surface forms the peculiar facets on the surface of the eye, which are known as the corneal lenses, while the cells of the epidermis give rise to lens-like bodies known as the crystalline cones.

The general features of the origin of the nervous system which have so far been made out by means of the study of embryology are the following:—

(1) That the nervous system of the higher Metazoa has been developed in the course of a long series of generations by a gradual process of differentiation of parts of the epidermis.

(2) That part of the central nervous system of many forms arose as a local collection of nerve-cells in the epidermis, in the neighbourhood of rudimentary organs of vision.

(3) That ganglion cells have been evolved from simple epithelial cells of the epidermis.

(4) That the primitive nerves were outgrowths of the original ganglion cells; and that the nerves of the higher forms are formed as outgrowths of the central nervous system.

The points on which embryology has not yet thrown a satisfactory light are:—

(1) The steps by which the protoplasmic processes, from the primitive epidermic cells, became united together so as to form a network of nerve-fibres, placing the various parts of the body in nervous communication.

(2) The process by which nerves became connected with muscles, so that a stimulus received by a nerve-cell could be communicated to and cause a contraction in a muscle.

Recent investigations on the anatomy of the Cœlenterata, especially of jelly-fish and sea-anemones, have thrown some light on these points, although there is left much that is still obscure.

The connection between this network and the muscular cells also probably took place by a process of contact and fusion.

Epithelial cells with muscular processes were discovered by Kleinenberg before epithelial cells with nervous processes were known, and he suggested that the epithelial part of such cells was a sense-organ, and that the connecting part between this and the contractile processes was a rudimentary nerve. This ingenious theory explained completely the fact of nerves being continuous with muscles; but on the further discoveries being made which I have just described, it became obvious that this theory would have to be abandoned, and that some other explanation would have to be given of the continuity between nerves and muscles. The hypothetical explanation just offered is that of fusion.

It seems very probable that many of the epithelial cells were originally provided with processes the protoplasm of which, like that of the Protozoa, carried on the functions of nerves and muscles at the same time, and that these processes united amongst themselves into a network. By a process of differentiation parts of this network may have become specially contractile, and other parts may have lost their contractility and become solely nervous. In this way the connection between nerves and muscles might be explained, and this hypothesis fits in very well with the condition of the neuro-muscular system as we find it in the Cœlenterata.

The nervous system of the higher Metazoa appears then to have originated from a differentiation of some of the superficial epithelial cells of the body, though it is possible that some parts of the system may have been formed by a differentiation of the alimentary epithelium. The cells of the epithelium were most likely at the same time contractile and sensory, and the differentiation of the nervous system may very probably have commenced, in the first instance, from a specialization in the function of part of a network formed of neuro-muscular prolongations of epithelial cells. A simultaneous differentiation of other parts of the network into muscular fibres may have led to the continuity at present obtaining between nerves and muscles.

Nerves, such as we find them in the higher types, originated from special differentiations of the nervous network, radiating from the parts of the central nervous system.

Such, briefly, is the present state of our knowledge as to the genesis of the nervous system. I ought not, however, to leave this subject without saying a few words as to the hypothetical views which the distinguished evolutionist Mr Herbert Spencer has put forward on this subject in his work on Psychology.

For Herbert Spencer nerves have originated, not as processes of epithelial cells, but from the passage of motion along the lines of least resistance. The nerves would seem, according to this view, to have been formed in any tissue from the continuous passage of nervous impulses through it. “A wave of molecular disturbance,” he says, “passing along a tract of mingled colloids closely allied in composition, and isomerically transforming the molecules of one of them, will be apt at the same time to form some new molecules of the same type,” and thus a nerve becomes established.

A nervous centre is formed, according to Herbert Spencer, at the point in the colloid in which nerves are generated, where a single nervous wave breaks up, and its parts diverge along various lines of least resistance. At such points some of the nerve-colloid will remain in an amorphous state, and as the wave of molecular motion will there be checked, it will tend to cause decompositions amongst the unarranged molecules. The decompositions must, he says, cause “additional molecular motion to be disengaged; so that along the outgoing lines there will be discharged an augmented wave. Thus there will arise at this point something having the character of a ganglion corpuscle.”

Although the present state of our knowledge on the genesis of the nervous system is a great advance on that of a few years ago, there is still much remaining to be done to make it complete.

The subject is well worth the attention of the morphologist, the physiologist, or even of the psychologist, and we must not remain satisfied by filling up the gaps in our knowledge by such hypotheses as I have been compelled to frame. New methods of research will probably be required to grapple with the problems that are still unsolved; but when we look back and survey what has been done in the past, there can be no reason for mistrusting our advance in the future.

"9. The lateral fins, as they were applied to support the body on the ground, became elongated, segmented, and narrowed, so that probably the line of the propterygium, or possibly that of the mesopterygium, became the cheiropterygial axis.

"10. The distal end of the incipient cheiropterygium either preserved and enlarged preexisting cartilages or developed fresh ones to serve fresh needs, and so grew into the developed cheiropterygium; but there is not yet enough evidence to determine what was the precise course of this transformation.

"11. The pelvic limb acquired a solid connection with the axial skeleton (a pelvic girdle) through its need of a point d'appui as a locomotive organ on land.

“12. The pelvic limb became also elongated; and when its function was quite similar to that of the pectoral limb, its structure became also quite similar (e.g. Ichthyosaurus, Plesiosaurus, Chelydra, &c.); but for the ordinary quadrupedal mode of progression it became segmented and inflected in a way generally parallel with, but (from its mode of use) in part inversely to, the inflections of the pectoral limb.”

“When we now detach the outermost phalanx from each side of the first horizontal zone, and with it the other phalanges of the same series, when we allow the remaining phalanges of this zone to coalesce into one piece (as, in nature, we find coalesced the carpals of Ceratodus and many phalanges in Selachian fins), and when we repeat this same process with the following zones and outer series, we arrive at an arrangement identical with what we actually find in Ceratodus.”

While the researches of Thacker and Mivart are strongly confirmatory of the view at which I had arrived with reference to the nature of the paired fins, other hypotheses as to the nature of the skeleton of the fins have been enunciated, both before and after the publication of my memoir, which are either directly or indirectly opposed to my view.

It will be seen that Huxley's idea of the primitive structure of the archipterygium is not easily reconcilable with the view that the paired fins are parts of a once continuous lateral fin, in that the skeleton of such a lateral fin, if it has existed, must necessarily have consisted of a series of parallel rays.

"All the varied forms which the skeleton of the free appendages exhibits may be derived from a ground-form which persists in a few cases only, and which represents the first, and consequently the lowest, stage of the skeleton in the fin—the archipterygium. This is made up of a stem which consists of jointed pieces of cartilage, which is articulated to the shoulder-girdle and is beset on either side with rays which are likewise jointed. In addition to the rays of the stem there are others which are directly attached to the limb-girdle.

“The metapterygium represents the stem of the archipterygium and the rays on it. The propterygium and the mesopterygium are evidently derived from the rays which still remain attached to the shoulder-girdle.”