The Works of Francis Balfour 1-19

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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

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This historic 1885 book edited by Foster and Sedgwick is the first of Francis Balfour's collected works published in four editions. Francis (Frank) Maitland Balfour, known as F. M. Balfour, (November 10, 1851 - July 19, 1882) was a British biologist who co-authored embryology textbooks.



Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
Modern Notes:

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Pages where the terms "Historic Textbook" and "Historic Embryology" appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms and interpretations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)


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Vol I. Separate Memoirs (1885)

XIX. Address to the Department of Anatomy and Physiology of The 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.

To attempt in the brief time which I have at my disposal to trace the influence of the Darwinian theory on all the branches of anatomy and physiology would be wholly impossible, and I shall confine myself to an attempt to do so for a small section only. There is perhaps no department of Biology which has been so revolutionised, if I may use the term, by the theory of animal evolution, as that of Development or Embryology. The reason of this is not far to seek. According to the Darwinian


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theory, the present order of the organic world has been caused by the action of two laws, known as the laws of heredity and of variation. The law of heredity is familiarly exemplified by the well-known fact that offspring resemble their parents. Not only, however, do the offspring belong to the same species as their parents, but they inherit the individual peculiarities of their parents. It is on this that the breeders of cattle depend, and it is a fact of every-day experience amongst ourselves. A further point with reference to heredity to which I must call your attention is the fact that the characters, which display themselves at some special period in the life of the parent, are acquired by the offspring at a corresponding period. Thus, in many birds the males have a special plumage in the adult state. The male offspring is not, however, born with the adult plumage, but only acquires it when it becomes adult.

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.

We are now in a position to follow out the consequences of these two laws in their bearing on development. Their application will best be made apparent by taking a concrete example. Let us suppose a spot on the surface of some very simple organism to become, at a certain period of life, pigmented, and therefore to be especially sensitive to light. In the offspring of this form, the pigment-spot will reappear at a corresponding period ; and there will therefore be a period in the life of the offspring during which there is no pigment-spot, and a second period in

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which there is one. If a naturalist were to study the life-history, or, in other words, the embryology of this form, this fact about the pigment-spot would come to his notice, and he would be justified, from the laws of heredity, in concluding that the species was descended from an ancestor without a pigment-spot, because a pigment-spot was absent in the young. Now, we may suppose the transparent layer of skin above the pigment-spot to become thickened, so as gradually to form a kind of lens, which would throw an image of external objects on the pigment-spot. In this way a rudimentary eye might be evolved out of the pigmentspot. A naturalist studying the embryology of the form with this eye would find that the pigment-spot was formed before the lens, and he would be justified in concluding, by the same process of reasoning as before, that the ancestors of the form he was studying first acquired a pigment-spot and then a lens. We may picture to ourselves a series of steps by which the simple eye, the origin of which I have traced, might become more complicated ; and it is easy to see how an embryologist studying the actual development of this complicated eye would be able to unravel the process of its evolution.

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.

A direct verification of this proposition is scarcely possible. There is ample ground for concluding that the forms from which


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existing animals are descended have in most instances perished ; and although there is no reason why they should not have been preserved in a fossil state, yet, owing to the imperfection of the geological record, palaeontology is not so often of service as might have been hoped.

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.


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Before the appearance of the Origin of Species many very valuable embryological investigations were made, but the facts discovered were to their authors merely so many ultimate facts, which admitted of being classified, but could not be explained. No explanation could be offered of why it is that animals, instead of developing in a simple and straightforward way, undergo in the course of their growth a series of complicated changes, during which they often acquire organs which have no function, and which, after remaining visible for a short time, disappear without leaving a trace.

No explanation, for instance, could be offered of why it is that a frog in the course of its growth has a stage in which it breathes like a fish, and then why it is like a newt with a long tail, which gradually becomes absorbed, and finally disappears. To the Darwinian the explanation of such facts is obvious. The stage when the tadpole breathes by gills is a repetition of the stage when the ancestors of the frog had not advanced in the scale of development beyond a fish, while the newt-like stage implies that the ancestors of the frog were at one time organized very much like the newts of to-day. The explanation of such facts has opened out to the embryologist quite a new series of problems. These problems may be divided into two main groups, technically known as those of phylogeny and those of organogeny. The problems of phylogeny deal with the genealogy of the animal kingdom. A complete genealogy would form what is known as a natural classification. To attempt to form such a classification has long been the aim of a large number of naturalists, and it has frequently been attempted without the aid of embryology. The statements made in the earlier part of my address clearly shew how great an assistance embryology is capable of giving in phylogeny ; and as a matter of fact embryology has been during the last few years very widely employed in all phylogenetic questions, and the results which have been arrived at have in many cases been very striking. To deal with these results in detail would lead me into too technical a department of my subject ; but I may point out that amongst the more striking of the results obtained entirely by embryological methods is the demonstration that the Vertebrata are not, as was nearly universally believed by older


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naturalists, separated by a wide gulf from the Invertebrata, but that there is a group of animals, known as the Ascidians, formerly united with the Invertebrata, which are now universally placed with the Vertebrata.

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.

In the first place every Metazoon commences its existence as a simple cell, in the sense above defined ; this cell is known as the ovum. The first developmental process which takes


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place consists in the division or segmentation of the single cell into a number of smaller cells. The cells then arrange themselves into two groups or layers known to embryologists as the primary germinal layers. These two layers are usually placed one within the other round a central cavity. The inner of the two is called the hypoblast, the outer the epiblast. The existence of these two layers in the embryos of vertebrated animals was made out early in the present century by Pander, and his observations were greatly extended by Von Baer and Remak. But it was supposed .that these layers were confined to vertebrated animals. In the year 1849, an d at greater length in 1859, Huxley demonstrated that the bodies of all the polype tribe or Coelenterata that is to say of the group to which the common polype, jelly-fish and the sea-anemone belong were composed of two layers of cells, and stated that in his opinion these two layers were homologous with the epiblast and hypoblast of vertebrate embryos. This very brilliant discovery came before its time. It fell upon barren ground, and for a long time bore no fruit In the year 1866 a young Russian naturalist named Kowalevsky began to study by special histological methods the development of a number of invertebrated forms of animals, and discovered that at an early stage of development the bodies of all these animals were divided into germinal layers like those in vertebrates. Biologists were not long in recognizing the importance of these discoveries, and they formed the basis of two remarkable essays, one by our own countryman, Professor Lankester, and the other by a distinguished German naturalist, Professor Haeckel, of Jena.

In these essays the attempt was made to shew that the stage in development already spoken of, in which the cells are arranged in the form of two layers enclosing a central cavity has an ancestral meaning, and that it is to be interpreted to signify that all the Metazoa are descended from an ancestor which had a more or less oval form, with a central digestive cavity provided with a single opening, serving both for the introduction of food and for the ejection of indigestible substances. The body of this ancestor was supposed to have been a double-walled sack formed of an inner layer, the hypoblast, lining the digestive


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cavity, and an outer layer, the epiblast. To this form Haeckel gave the name of gastraea or gastrula.

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 gastraea 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 Ccelenterata, 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.

We have seen that the epiblast represents the skin or epidermis of the simple sack-like ancestor common to all the Metazoa. In all the higher Metazoa it gives rise, as might be expected, to the epidermis, but it gives rise at the same time to a number of other organs ; and, in accordance with the principles laid down in the earlier part of my address, it is to be concluded that the organs so derived have been formed as differentiations of


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the primitive epidermis. One of the most interesting of recent embryological discoveries is the fact that the nervous system is, in all but a very few doubtful cases, derived from the epiblast. This fact was made out for vertebrate animals by the great embryologist Von Baer; and the Russian naturalist Kowalevsky, to whose researches I have already alluded, shewed that this was true for a large number of invertebrate animals. The derivation of the nervous system from the epiblast has since been made out for a sufficient number of forms satisfactorily to establish the generalization that it is all but universally derived from the epiblast.

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 Coelenterata, fills up some of the gaps left from our study of embryology.


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From embryology we learn that the ganglion-cells of the central part of the nervous system are originally derived from the simple undifferentiated epithelial cells of the surface of the body. We further learn that the nerves are out-growths of the central nervous system. It was supposed till quite recently that the nerves in Vertebrates were derived from parts of the middle germinal layer or mesoblast, and that they only became secondarily connected with the central nervous system. This is now known not to be the case, but the nerves are formed as processes growing out from the central part of the nervous system.

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-cesophageal 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-cesophageal 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-cesophageal 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.

It would be easy to quote further instances of the same kind, but I trust that the two which I have given will be sufficient to shew the kind of relation which often exists between the organs


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of special sense, especially those of vision, and the central nervous system. It might have been anticipated a priori that organs of special sense would only appear in animals provided with a well-developed central nervous system. This, however, is not the case. Special cells, with long delicate hairs, which are undoubtedly highly sensitive structures, are present in animals in which as yet nothing has been found which could be called a central nervous system ; and there is every reason to think that the organs of special sense originated pari passn with the central nervous system. It is probable that in the simplest organisms the whole body is sensitive to light, but that with the appearance of pigment-cells in certain parts of the body, the sensitiveness to light became localised to the areas where the pigment-cells were present. Since, however, it was necessary that stimuli received by such organs should be communicated to other parts of the body, some of the epidermic cells in the neighbourhood of the pigment-spots, which were at first only sensitive, in the same manner as other cells of the epidermis, became gradually differentiated into special nerve-cells. As to the details of this differentiation, embryology does not as yet throw any great light ; but from the study of comparative anatomy there are grounds for thinking that it was somewhat as follows : Cells placed on the surface sent protoplasmic processes of a nervous nature inwards, which came into connection with nervous processes from similar cells placed in other parts of the body. The cells with such processes then became removed from the surface, forming a deeper layer of the epidermis below the sensitive cells of the organ of vision. With these cells they remained connected by protoplasmic filaments, and thus they came to form a thickening of the epidermis underneath the organ of vision, the cells of which received their stimuli from those of the organ of vision, and transmitted the stimuli so received to other parts of the body. Such a thickening would obviously be the rudiment of a central nervous system, and it is easy to see by what steps it might become gradually larger and more important, and might gradually travel inwards, remaining connected with the sense organ at the surface by protoplasmic filaments, which would then constitute nerves. The rudimentary eye would at first merely consist partly of cells sensitive to light, and partly of optical structures


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constituting the lens, which would throw an image of external objects upon it, and so convert the whole structure into a true organ of vision. It has thus come about that, in the development of the individual, the retina or sensitive part of the eye is first formed in connection with the central nervous system, while the lenses of the eye are independently evolved from the epidermis at a later period.

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

In our own country Mr Romaines has conducted some interesting physiological experiments on these forms ; and Professor Schafer has made some important histological investigations upon them. In Germany a series of interesting researches have also been made on them by Professors Kleinenberg, Claus and


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Eimer, and more especially by the brothers Hertwig, of Jena. Careful histological investigations, especially those of the lastnamed authors, have made us acquainted with the forms of some very primitive types of nervous system. In the common sea-anemones there are, for instance, no organs of special sense, and no definite central nervous system. There are, however, scattered throughout the skin, and also throughout the lining of the digestive tract, a number of specially modified epithelial cells, which are no doubt delicate organs of sense. They are provided at their free extremity with a long hair, and are prolonged on their inner side into a fine process which penetrates the deeper part of the epithelial layer of the skin or digestive wall. They eventually join a fine network of protoplasmic fibres' which forms a special layer immediately within the epithelium. The fibres of this network are no doubt essentially nervous. In addition to fibres there are, moreover, present in the network cells of the same character as the multipolar ganglion-cells in the nervous system of Vertebrates, and some of these cells are characterized by sending a process into the superjacent epithelium. Such cells are obviously epithelial cells in the act of becoming nerve-cells ; and it is probable that the nerve-cells are, in fact, sense-cells which have travelled inwards and lost their epithelial character.

There is every reason to think that the network just described is not only continuous with the sense-cells in the epithelium, but that it is also continuous with epithelial cells which are provided with muscular prolongations. The nervous system thus consists of a network of protoplasmic fibres, continuous on the one hand with sense-cells in the epithelium, and on the other with muscular cells. The nervous network is generally distributed both beneath the epithelium of the skin and that of the digestive tract, but is especially concentrated in the disc-like region between the mouth and tentacles. The above observations have thrown a very clear light on the characters of the nervous system at an early stage of its evolution, but they leave unanswered the questions (i) how the nervous network first arose, and (2) how its fibres became continuous with muscles. It is probable that the nervous network took its origin from processes of the sense-cells. The processes of the different cells probably first met and then fused


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together, and, becoming more arborescent, finally gave rise to a complicated network.

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 Ccelenterata.

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.


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Local differentiations of the nervous network, which was no doubt distributed over the whole body, took place on the formation of organs of special sense, and such differentiations gave rise to the formation of a central nervous system. The central nervous system was at first continuous with the epidermis, but became separated from it and travelled inwards. Ganglion-cells took their origin from sensory epithelial cells, provided with prolongations, continuous with the nervous network. Such epithelial cells gradually lost their epithelial character, and finally became completely detached from the epidermis.

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."


AND PHYSIOLOGY OF THE BRITISH ASSOCIATION. 713

These hypotheses of Herbert Spencer, which have been widely adopted in this country, are, it appears to me, not borne out by the discoveries to which I have called your attention to-day. The discovery that nerves have been developed from processes of epithelial cells, gives a very different conception of their genesis to that of Herbert Spencer, which makes them originate from the passage of nervous impulses through a tract of mingled colloids ; while the demonstration that ganglion-cells arose as epithelial cells of special sense, which have travelled inwards from the surface, admits still less of a reconciliation with Herbert Spencer's view on the same subject.

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.