Form and Function Part 20

We have seen this physiological att.i.tude expressed with the utmost clearness by the founder of the cell-theory himself; we shall see the same att.i.tude taken up by most of his successors. Thus Vogt, who was later to become one of the protagonists of materialism in Germany, developed in his memoir on the embryology of _Coregonus_[293] the theory of the independent or individual life of the cell. "Each cell," he wrote, "represents in some measure a separate organism, and while their development necessarily conforms to the general plan and the particular tendencies of the parent organism, they nevertheless each follow their own particular tendency and do not lose their independence until, by reason of the metamorphoses which they undergo, they lose their cellular nature" (p. 275).

And again, "... we are obliged to admit the existence in the cell of an independent life, which makes its development self-sufficient.... Each cell consequently represents a little independent organism, which a.s.similates foreign substances, builds them up, and rejects those that are useless; from this point of view the embryo can be compared up to a certain point with a zoophyte stock, of which each polyp, while living its own independent life, is yet incorporated in the common corm, which impresses its distinctive character upon every polyp" (p. 293).

Cla.s.sical expression was given to the "colonial theory" of the organism by Virchow in his lectures on "Cellular Pathology."[294] For Virchow the organism resolves itself into an a.s.semblage of living centres, the cells; the organism has no real existence as a unity, for there is no one single centre from which its activities are ruled. Even the nervous system, which appears to act as a co-ordinating centre, is itself an aggregate of discrete cells. "A tree is a body of definite and orderly composition, the ultimate elements of which, in every part of it, in leaf and root, in stem and flower, are cellular elements--so also are animal forms. _Every animal is a sum of vital units_, each of which possesses the full characteristics of life. The character and the unity of life cannot be found in one definite point of a higher organisation, for example in the brain of man, but only in the definite, constantly recurring disposition shown individually by each single element. It follows that the composition of the major organism, the so-called individual, must be likened to a kind of social arrangement or society, in which a number of separate existences are dependent upon one another, in such a way, however, that each element possesses its own particular activity, and, although receiving the stimulus to activity from the other elements, carries out its own task by its own powers" (2nd ed., pp. 12-13).

a.n.a.lysis, decomposition, or disintegration of the organism is here pushed to its extreme point, and the problem of recomposition, synthesis and co-ordination shirked or forgotten.

The harmful influence of the cell-theory upon morphology did not pa.s.s unnoticed by the broader-minded zoologists of the day. Virchow's earlier paper[295] on the application of the cell-theory to physiology and pathology called forth a vigorous protest from Reichert,[296] who discussed in a very instructive way the contrast between the older "systematic" and the newer "atomistic" att.i.tude to living Nature.

Is it really true, he asks, that the cell is the dominant element in all organisation; is the cell comparable in importance to the atom of the chemists; or is it not rather the servant of a higher regulatory power?

Johannes Muller, who was Reichert's master, had in his _Physiology_[297]

argued splendidly for the existence of a creative force which guides and rules development, and brings to pa.s.s that unity and harmony of composition which distinguish living things from inorganic products.

Reichert sought in vain in the writings of the biological "atomists" for any smallest recognition of these broader characteristics of living things upon which Muller had rightly laid stress. For the atomists the cell was the only element of form; they ignored the combination of cells to form tissues, of tissues to form organs, of organs to form an organism. For the morphologists the cell was one element among many, and the lowest of all.

The difference of att.i.tude is clearly shown if we consider from the two points of view a complicated organ-system such as the central nervous system. The atomist sees in this a mere aggregate of cells or at the most of groups of cells. "The morphologist," on the other hand, "sees in the central nervous system a _proximate_ element in the composition of the body--a primitive organ. From this point of view he apprehends and judges its morphological relations with, in the first place, the other co-ordinated primitive organs in the system as a whole; in all this the cells remain in the background, and have nothing to do directly with the determination of these morphological relations" (p. 6). Within the nervous system there are separate organs which stand to one another in a definite morphological and functional relationship. These organs are, it is true, composed of cells; but between the form and connections of these organs and the cells which compose them there is no direct and necessary relation (p. 6). It is true that the cell is the ultimate element of organic form, and that all development takes place by multiplication and form-change of cells. Yet is the cell in all this not independent of the unity of the developing embryo, and what the cells produce, they produce, so to speak, not of their own free will, nor by chance, but under the guiding influence of the unity of the whole, and in a certain measure as its agents (p. 7). The atomists will not admit the truth of this; they see in development nothing more than a process of the form-change and multiplication of cells. The full meaning of development escapes them, for they take no cognisance of the increasing complexity of the embryo, of the separating-out of tissues, of the moulding of organs, of the harmonious adaptation and adjustment of the parts to form a working whole.

In general, the fault of the atomists is that they do not respect the limits which Nature herself has prescribed to the process of logical a.n.a.lysis and disintegration of the organism; they do not recognise the existence of natural and rational units or unities; they forget the one great principle of rational a.n.a.lysis, "that, by universally valid, inductive, logical method, natural objects must in all cases be accepted and dealt with in the combination and concatenation in which they are given" (p. 10).

The atomists at least recognised one natural organic element, the cell; the materialistic physiologists of the time resolved even this unity into an aggregate of inorganic compounds, and regarded the organism itself as nothing but a vastly complicated physico-chemical mechanism.

From this point of view morphology had no right of existence, and we find Ludwig, one of the foremost of the materialistic school, maintaining that morphology was of no scientific importance, that it was nothing more than an artistic game, interesting enough, but completely superseded and robbed of all value by the advance of materialistic physiology.[298]

Naturally enough, morphologists did not accept this rather contemptuous estimate of their science, but held firmly to the morphological att.i.tude. So Leuckart in his reply to Ludwig, so Rathke in a letter to Leuckart published in that reply, so Reichert in his _Bericht_, so J. V.

Carus in his _System der thierischen Morphologie_,[299] upheld the validity, the independence, of morphological methods. Leuckart and Rathke called attention to the absolute impossibility of explaining by materialistic physiology the unity of plan underlying the diversity of animal form. J. V. Carus, who was convinced of the validity of physiological methods within their proper sphere, drew a sharp distinction between systematics and morphology on the one hand, and physiology on the other. Physiology had nothing to do with the problems of form at all; its business was to study the physical and chemical processes which lay at the base of all vital activities. Morphology, on its part, had to accept form as something given, and to study the abstract relations of forms to one another. "On this point," he writes, "stress is to be laid, that morphology has to do with animal form as something _given_ by Nature, that though it follows out the changes taking place during the development of an animal and tries to explain them, it does not enquire after the conditions whose necessary and physical consequence this form actually is" (p. 24). He expressed indeed a pious hope (p. 25) that physiology might one day be so far advanced that it could attempt with some hope of success to discover the physico-chemical determinism of form, but this remained with him merely a pious hope. Reichert, in his _Bericht_, applied to the rather wild theorisings of the physiologist Ludwig the same clear commonsense criticism that he bestowed on the other "atomists."

It would take too long to describe the great development that materialistic physiology took at this time, and to show how the separation of morphology from physiology, which originally took place away back in the 17th century, had by this time become almost absolute.

The years towards the end of the first half of the century marked indeed the beginning of the cla.s.sical period as well of physiology as of dogmatic materialism. Moleschott and Buchner popularised materialism in Germany in the 'fifties, while Ludwig, du Bois Reymond and von Helmholtz began to apply the methods of physics to physiology. In France, Claude Bernard was at the height of his activity, rivalled by workers almost as great. The doctrine of the conservation of energy was established about this same time.

Between the cell-theory on the one side, and physiology on the other, it was a wonder that morphology kept alive at all. The only thing that preserved it was the return to the sound Cuvierian tradition which had been made by many zoologists in the 'thirties and 'forties. It is a significant fact that this return to the functional att.i.tude coincided in the main with the rise of marine zoology, and that the man who most typically preserved the Cuvierian att.i.tude, H. Milne-Edwards, was also one of the first and most consistent of marine biologists. Milne-Edwards describes in his interesting _Rapport sur les Progres recents des Sciences zoologiques en France_ (Paris) 1867, how "About the year 1826, two young naturalists, formed in the schools of Cuvier, Geoffroy and Majendie, considered that zoology, after having been purely descriptive or systematic and then anatomical, ought to take on a more physiological character; they considered that it was not enough to observe living objects in the repose of death, and that it was desirable to get to understand the organism in action, especially when the structure of these animals was so different from that of man that the notions acquired as to the special physiology of man could not properly be applied to them" (p. 17). The two young naturalists were H.

Milne-Edwards and V. Audouin. In pursuance of these excellent ideas they set to work to study the animals of the seash.o.r.e, producing in 1832-4 two volumes of _Recherches pour servir a l'histoire naturelle du littoral de la France_. After Audouin's early death A. de Quatref.a.ges was a.s.sociated with Milne-Edwards in this pioneer work, and their valiant struggles with insufficient equipment and lack of all laboratory accommodation, and the rich harvest they reaped, may be read of in Quatref.a.ge's fascinating account of their journeyings.[300] Note that though they called themselves physiologists they meant by physiology something very different from the mere physical and chemical study of living things. They were interested, as Cuvier was, primarily in the problems of form; they sought to penetrate the relation between form and function; their chief aim was, therefore, the study not of physiology[301]

in the restricted sense, but physiological morphology. As a matter of fact they produced more taxanomic and anatomical work than work on physiological morphology, but this was only natural, since such a wealth of new forms was disclosed to their gaze. Milne-Edwards' masterly _Histoire Naturelle des Crustaces_[302] and A. de Quatref.a.ge's _Histoire Naturelle des Anneles marins et d'eau douce_[303] were typical products of their activity.

In the North, men like Sars and Loven were starting to work on the littoral fauna of the fjords; in Britain, Edward Forbes was opening up new worlds by the use of the dredge; Johannes Muller was using the tow-net to gather material for his masterly papers on the metamorphoses of Echinoderms.[304] Work on the taxonomy and anatomy of marine animals was in general in full swing by the 'fifties and 'sixties.

This return to Nature and to the sea had a very beneficial effect upon morphology, bringing it out from the laboratory to the open air and the seash.o.r.e. It saved morphology from formalism and aridity, and in particular from a certain narrowness of outlook born of too close attention paid to the details of microscopical anatomy. It brought morphologists face to face again with the wonderful diversity of organic forms, with the unity of plan underlying that diversity, with the admirable adjustment of organ to function and of both to the life of the whole.

Milne-Edwards' theoretical views, as expounded in his _Introduction a la zoologie generale_ (1851), well reflect this Cuvierian att.i.tude.[305] He acknowledges himself the debt he owes to Cuvier; "the further I advance in the study of the sciences which he cultivated with so sure a hand,"

he writes in 1867, "the more I venerate him."

Milne-Edwards frankly takes up the teleological standpoint, and interprets organic forms on the a.s.sumption that they are purposive and rationally constructed. "To arrive at an understanding of the harmony of the organic creation," he writes, "it seemed to me that it would be well to accept the hypothesis that Nature has gone about her work as we would do ourselves according to the light of our own intelligence, if it were given us to produce a similar result. Comparing and studying living things as if they were machines created by the industry of man, I have tried to grasp the manner in which they might have been invented, and the principles whose application would have led to the production of such an a.s.semblage of diversified instruments" (p. 435). The problem is to discover the laws which rule the diversity of organic forms. The first and most obvious of these laws is the "law of economy," or the law of unity of type. Nature, as Cuvier pointed out, has not had recourse to all the possible forms and combinations of organs; she appears to work with a limited number of types and to get the greatest possible diversity out of these by varying the proportions of the const.i.tutive materials of structure. Within the limits of each type Nature has brought about diversity by raising her creatures to different degrees of perfection. This is the second law of organic form, and it is this law that Milne-Edwards chiefly elaborates. Degrees of perfection mean for him, as for Aristotle, primarily degrees of perfection of function, but since structure is necessarily in close relation with function, perfection of function brings in its train increased perfection of organisation. This can only be attained by a division of labour[306] among the organs and by their consequent differentiation. An animal is like a workshop where some complicated product is manufactured, and the organs are like the workmen. Each workman has his own special piece of work to do, at which he becomes thoroughly expert; and the finished product is manufactured more rapidly and efficiently by the co-operation of workers each skilled in one department than it would be if each workman had to produce the whole. Applied to the organism this principle of the division of labour means the differentiating out of the separate functions, their localisation in different parts of the organism, and their co-ordination to produce a combined result.

This differentiation of functions implies a corresponding differentiation of organs, but it is functional differentiation which always takes the lead. "Where division of labour has not been introduced into the organism there must exist a great simplicity of structure. But just as uniformity in the functions of the different parts of the body implies a uniformity in their mode of const.i.tution, so diversity in function must be accompanied by particularities in structure; and, in consequence also, the number of dissimilar parts must be augmented and the complication of the machine increased" (p. 463). Since function comes before form there is not always a special organ for every function. "It is a grave error to believe that a particular function can be performed only by one and the same organ. Nature can arrive at the desired result by various ways, and when we look down through the animal kingdom from the highest to the lowest forms we see that the function does not disappear even when the special instrument provided for the purpose in the higher types ceases to exist" (p 470).

Nature, holding fast to the law of economy, does not even always create a new organ for a new function; she may simply adapt an undifferentiated part to special functions, or she may even convert to other uses an organ already specialised (p. 464). So, for example, the function of respiration is in the lowest animals diffused indifferently over the whole surface of the body, and only as organisation advances is it localised in special organs, such as gills. Now suppose that Nature wishes to adapt a fish, which breathes by gills, to life in the air; she does not create an organ specially for this purpose, but utilises the moist gill-chamber (_e.g._, in _Anabas scandens_), modifying it in certain ways so that the fish can take advantage of the oxygen it contains. But this gill-chamber lung is at best a makeshift, and when she comes to the more definitely terrestrial Amphibia Nature gives up the attempt to use the gill-chamber as a lung, and creates a new organ, the true vertebrate lung, specially adapted for breathing air (p. 475).

But whatever means Nature adopts, her aim is always the same--to specialise, to differentiate, to produce diversity from uniformity.

Differentiation not only raises the level of organisation; it usually also takes the direction of adaptation to particular habits of life, and this is perhaps the most fruitful cause of diversity. Everywhere we find animals specialised in adaptation to their environment--to life in air or water, or on land--and many of their most striking differences are due to this cause. But adaptation may also act in reducing diversity, for there necessarily occur many instances of parallel adaptation or convergence. So we get the extraordinary parallelism between the families of marsupials and the orders of placentals,[307] the remarkable similarity between the respiratory organs of land-crabs and air-breathing fish--to mention only two out of an immense range of a.n.a.logous facts.

The last cause of diversity that Milne-Edwards adduces is what he calls a "borrowing" of peculiarities of structure from another systematic group. Thus, "among reptiles, the tortoises seem to have borrowed from birds some of their characteristic features of organisation; and among the sauroid fishes the piscine type seems to have been influenced by the type from which reptiles are derived" (p. 479). So many riddles that, a little later on, stimulated the ingenuity of the evolutionists!

Such, then, were the factors which Milne-Edwards considered adequate to explain the rich variety of animal forms. We cannot do better than quote his own summary of his doctrine:--"To sum up, then, the great differences introduced by Nature into the const.i.tution of animals seem to depend essentially upon the existence of a certain number of general plans or distinct types, upon the perfecting in various degrees either of the whole or of parts of each of these structural plans, upon the adaptation of each type to varied conditions of existence, and upon the secondary imitation of foreign types by certain derivatives of each particular type" (p. 480).

We have laid stress on the fact that Milne-Edwards put function before form, for this is the mark of the true Cuvierian. With it goes the belief that Nature forms new parts to meet new requirements, that she is not limited, as Geoffroy thought, to a definite number of "materials of organisation," but can produce others at need. Cuvier held, for example, that many of the muscles and even the bones of fish were peculiar to them, and without h.o.m.ologues in the other Vertebrates, having been created by Nature for special ends.[308] So, too, Johannes Muller, who in many ways and not least in his sane vitalism was a follower of the Cuvierian tradition, recognised that many of the complicated cartilages in the skull of Cyclostomes were specially formed for the important function of sucking, and had no equivalent in other fish.[309]

So, too, the embryologists after Cuvier often came across instances of the special formation of parts to meet temporary needs. Thus Reichert interpreted the "palatine" and "pterygoid," which are formed in the mouth of the newt larva by a fusion of conical teeth, as special adaptations to enable the little larva to lead a carnivorous life.[310]

Not many years after the publication of Milne-Edwards' _Introduction a la zoologie generale_ (1851) there appeared a book by H. G. Bronn in which was offered a very similar a.n.a.lysis of organic diversity. The curious thing was that Bronn approached the problem from quite a different standpoint, from the standpoint, indeed, of _Naturphilosophie_. Of this the t.i.tle of the book is itself sufficient proof--_Morphologische Studien uber die Gestaltungs-gesetze der Naturkorper uberhaupt und der organischen insbesondere_ (Leipzig and Heidelberg, 1858).[311] The linking up of organic with inorganic form is characteristic; there is much talk, too, in the book of _Urstoffe_ and _Urkrafte_, but underlying the _Naturphilosophie_ we can trace the same Cuvierian treatment of form, and see crystallise out laws of progressive development that bear no small a.n.a.logy with the laws established by Milne-Edwards.

According to Bronn, the ideal fundamental form of the plant is an ovoid or strobiloid[312] body, for a plant reaches out in two directions in search of food--towards the sun and towards the earth. Animals differ from plants in being endowed with sensation and mobility (_cf._ Aristotle and Cuvier), and it is this characteristic that gives them their distinctive form. The main types of animal form--the Amorphozoa, Actinozoa, and Hemisphenozoa--are essentially adaptations to particular modes of locomotion. Animals either are fixed, or they move in all directions without reference to any definite axis, or they move in one main direction.

The Amorphozoa or shapeless animals include many of the Protozoa and sponges; they have no typical form, and most of them are sessile. The Actinozoa include such animals as the Coelentera, which are fixed, and the Echinoderms, which have a central point and move indifferently along any radial axis; their form differs from the strobiloid mainly in having radiate rather than spiral symmetry. The Hemisphenozoa, or bilaterally symmetrical animals, include all those that habitually move forward; they have a front end and a hind end, a dorsal surface and a ventral, and the mouth, sense-organs and "brain" are concentrated in the front end to form a head--all in direct adaptation to this forward movement; they make up the vast majority of animals.

The fundamental forms of living things are, however, merely so many themes on which a mult.i.tude of further variations are woven, through the action of the laws which rule the detail of organic diversities. These further laws may be set down under four main heads. Under the first comes the law of the existence of certain fundamentally distinct structural types, which are distinguished from one another by their ground-form, by the number of organ-systems, and by the number of h.o.m.otypic organs they possess, but princ.i.p.ally by the relative position of the organs to one another (principle of connections). The form and connections of the nervous system are of particular importance in distinguishing the types (_cf._ Cuvier). The second factor in the diversity of organic form is the action of certain laws of progressive development[313] (_Entwickelungsgesetze_), which bear the same relation to the development of the animal kingdom as the laws of individual development bear to the development of the embryo, for organs appear in the different animal series in much the same order and manner as they develop in the individual. These laws are (1) progressive differentiation of functions and organs; (2) numerical reduction of serially repeated parts; (3) concentration of functions and their organs in particular parts of the body; (4) centralisation of organ-systems and parts of such, so that they come to depend upon one central organ; (5) internalisation of the "n.o.blest" organs, unless these are necessarily external, and (6) increase in size of the whole or of parts. Of these the law of differentiation is by far the most important, and most of the others are in a sense merely special cases of this fundamental law. To this law of differentiation is due the increase in complexity or perfection of organisation which is shown by all the animal series.

Bronn himself recognised the great similarity of this law of progressive differentiation to Milne-Edwards' principle of the division of labour; he seems, however, to have arrived at it independently.

Bronn's third factor in the production of variety of form is adaptation to environment, or better, functional response to environment. Bronn gives an excellent account of adaptational modifications and calls attention, just as Milne-Edwards did, to the numerous a.n.a.logies of structure which adaptation brings about. He works out the interesting view that there is some connection between cla.s.sificatory groups and adaptational forms, especially such as are connected with the function of locomotion:--"Based upon a common characteristic method of locomotion are whole or nearly whole sub-phyla (Hexapoda), cla.s.ses (mammals and reptiles, birds, fishes, gastropods, pteropods, brachiopods, Bryozoa, Rotifera, jelly-fish, polypes, sponges), sub-cla.s.ses (mobile and immobile lamellibranchs, echinoderms, walking and swimming Crustacea, parasitic and free-living worms, and so on), often, however, only orders and quite small groups (snakes, eels, bats, sepias, medusae, etc.)" (p.

141).

It was characteristic of the 'forties and 'fifties that transcendental anatomy, along with Nature-philosophy, went rather out of fashion, its false simplicities and premature generalisations being overwhelmed by the flood of new discoveries. A few stalwarts indeed upheld transcendental views. We have already discussed the morphological system built up by Richard Owen in the late 'forties, a system transcendental in its main lines. We have seen the vertebral theory of the skull still maintained in the 'fifties by such men as Reichert and Kolliker, and we find J. V. Carus in 1853[314] taking it as almost conclusively proved.[315]

We may mention, too, as showing clear marks of the influence of transcendental ideas, L. Aga.s.siz's work on the principles of cla.s.sification.[316] And Serres, who was Geoffroy's chief disciple, recanted not a whit of his doctrine of recapitulation, but re-affirmed and expanded it from time to time, and particularly in a lengthy memoir published in 1860.[317] But in general we may say that pure morphology in the Geoffroyan or Okenian sense was becoming gradually discredited. A curious indication of this is seen in the fact that not only the idea but the very word "Archetype" came to be regarded with suspicion. Thus even J. V. Carus, who had much affinity with the transcendentalists, wrote of the vertebrate archetype (which he took over almost bodily from Owen)--"It may here be observed that this schema may be used as a methodological help, but it is not to be placed in the foreground"

(_loc. cit._, p. 395). Huxley, who was definitely a follower of von Baer, was much more outspoken with regard to ideal types. In an important memoir on the general anatomy of the Gastropoda and Cephalopoda,[318] he set himself the task of reducing all their complex forms to one type. In summing up, he writes:--"From all that has been stated, I think that it is now possible to form a notion of the archetype of the Cephalous Mollusca, and I beg it to be understood that in using this term, I make no reference to any real or imaginary 'ideas'

upon which animal forms are modelled. All that I mean is the conception of a form embodying the most general propositions that can be affirmed respecting the Cephalous Mollusca, standing in the same relation to them as the diagram to a geometrical theorem, and like it, at once imaginary and true" (i., p. 176). Again, in his Croonian lecture on the theory of the vertebrate skull, he remarks that a general diagram of the skull could easily be given. "There is no harm," he continues, "in calling such a convenient diagram the 'Archetype' of the skull, but I prefer to avoid a word whose connotation is so fundamentally opposed to the spirit of modern science" (_Sci. Memoirs_, vol. i., p. 571).

It is instructive to find that between Serres and Milne-Edwards there existed the same antagonism as between von Baer and the German transcendentalists. Milne-Edwards was a constant critic of the law of parallelism which Serres continued to uphold with little modification for over thirty years, just as von Baer was a critic of that form of the doctrine which was current in the early part of the century. As early as 1833, Milne-Edwards, through his studies of crustacean development,[319]

had come to the conclusion, independently of von Baer, that development always proceeded from the general to the special; that cla.s.s characters appeared before family characters, generic characters before specific.

In an interesting paper published in 1844,[320] he discussed the relation of this law of development to the problems of cla.s.sification, and arrived at results almost identical with those set forth by von Baer in his Fifth Scholion.

Like von Baer he rejected completely the theory of parallelism and the doctrine of the scale of beings; like von Baer he held that the type of organisation--of which there are several--is manifested in the very earliest stages and becomes increasingly specialised throughout the course of further development; like von Baer, too, he sketched a cla.s.sification based upon embryological characters.

These views were further developed in his volume of 1851, and also in his _Rapport_ of 1867.

They brought him into conflict with his confrere in the Academy of Sciences, etienne Serres, who in a number of papers published in the 'thirties and 'forties,[321] and particularly in his comprehensive memoir of 1860, still maintained the theory of parallelism and the doctrine of the absolute unity of type. His memoir of 1860 shows how completely Serres was under the domination of transcendental ideas. Much of it indeed goes back to Oken. "The animal kingdom," he writes, "may be considered in its entirety as a single ideal and complex being" (p.

141). His views have become a little more complicated since his first exposition of them in 1827, and he has been forced to modify in some respects the rigour of his doctrine. But he still holds fast to the main thesis of transcendentalism--the absolute unity of plan of all animals, vertebrate and invertebrate alike,[322] the gradual perfecting of organisation from monad to man, the repet.i.tion in the embryogeny of the higher animals of the "zoogeny" of the lower.

He recognised, however, that the idea of a simple scale of beings is only an abstraction, and that the true repet.i.tion is of organs rather than of organisms. He was willing even to admit, at least in the later pages of his memoir, that there might be not one animal series but several parallel series, as had been suggested by Isidore Geoffroy St Hilaire (p. 749). In general, his views are now less dogmatic than they were in his earlier writings, but they are not for all that changed in any essential. For, in summing up his main results, he writes, "The whole animal kingdom can in some measure be regarded ideally as a single animal, which, in the course of formation and metamorphosis in its diverse manifestations, here and there arrests its own development, and thus determines at each point of interruption, by the very state it has reached, the distinctive characters of the phyla, the cla.s.ses, families, genera, and species" (p. 833).[323]

To settle the dispute pending between two of its most ill.u.s.trious members, the Academy proposed in 1853, as the subject of one of its prizes, "the positive determination of the resemblances and differences in the comparative development of Vertebrates and Invertebrates." A memoir was presented the next year by Lereboullet[324] which met with the approval of the Academy in so far as its statements of fact were concerned, but seemed to them to require amplification in its theoretical part. But even in this memoir Lereboullet was able to show that the balance of evidence was greatly in favour of Milne-Edwards'

views, and his general conclusions in 1854 were that "in the presence of such fundamental differences, one is obliged to give up the idea of one single plan in the formation of animals; while, on the contrary, the existence of diverse plans or types is clearly demonstrated by all the facts" (p. 79). To fulfil the Academy's requirements, Lereboullet continued his work, and in 1861-63 he published a series of elaborate monographs[325] on the embryology of the trout, the lizard and the pond-snail _Lymnaea_, and rounded off his work with a full discussion[326]

of the theoretical questions involved. In this considered and authoritative judgment he completely disposed of Serres' theories of the unity of plan and the unity of genetic formation. Except in the very earliest stages of oogenesis there is no real similarity between the development of a Zoophyte, a Mollusc, an Articulate and a Vertebrate, but each is stamped from the beginning with the characteristics of its type. The lower animals are not, and cannot possibly be the permanent embryos of the higher animals. "The results which I have obtained," he writes, "are diametrically opposed to the theory of the zoological series const.i.tuted by stages of increasing perfection, a theory which tries to demonstrate in the embryonic phases of the higher animals a repet.i.tion of the forms which characterise the lower animals, and which has led to the a.s.sertion that the latter are permanent embryos of the former. The embryo of a Vertebrate shows the vertebrate type from the very beginning, and retains this type throughout the whole course of its development; it never is, and never can be, either a Mollusc or an Articulate" (xx., p. 54).

"We are led to establish ... as the general result of our researches, the existence of several types, and, consequently, of different plans, in the development of animals. These different types are manifested from the very beginning of embryonic life; the characters distinguishing them are therefore primordial, and we can say with M. Milne-Edwards that _everything goes to prove that the distinction established by Nature between animals belonging to different phyla is a primordial distinction_" (p. 58).

In other directions also von Baer's work was confirmed and extended by later observers--those parts of it particularly that had reference to the germ-layer theory, and to the concept of histological differentiation. His germ-layer theory was accepted in its main lines by Rathke, Bischoff and Lereboullet, and applied by them to the mult.i.tude of new facts they discovered. Rathke, in particular, was a firm upholder of the doctrine, and made considerable use of it in his writings.[327]

Even before the publication of von Baer's book he had interpreted in terms of the germ-layer theory sketched by his friend Pander the splitting of the blastoderm which occurs in the early development of _Astacus_, whereby there are formed a serous and a mucous layer, one inside the other--like the coats of an onion, to use his own expressive phrase.[328]

An ingenious application of the Pander-Baer theory was made by Huxley, who compared the outer and inner cell-layers which form the groundwork of the Coelentera with the serous and mucous layers of the vertebrate germ.[329] He laid stress, it is true, rather on the physiological than on the morphological resemblance. "A complete ident.i.ty of structure," he writes, "connects the 'foundation membranes' of the Medusae with the corresponding organs in the rest of the series; and it is curious to remark, that throughout, the outer and inner membranes appear to bear the same physiological relation to one another as do the serous and mucous layers of the germ; the outer becoming developed into the muscular system, and giving rise to the organs of offence and defence; the inner, on the other hand, appearing to be more closely subservient to the purposes of nutrition and generation" (p. 24). Von Baer had already hinted at this h.o.m.ology in the second volume of his _Entwickelungsgeschichte_ (1837), where he says with reference to the separation of the blastoderm of the chick into two layers. "Yet originally there are not two distinct or even separable layers, it is rather the two surfaces of the germ which show this differentiation, just as polyps show the same contrast of an external surface and an internal digestive surface. In between the two layers there is in our germ as in the polyp an indifferent ma.s.s" (p. 67). The terms ectoderm and entoderm were introduced by Allman[330] in 1853 for the two cell-layers in the Hydrozoa.

Remak is the second great name in the history of the germ-layer theory.