The attempt was an interesting one, but most morphologists wisely adhered to the old concept of h.o.m.ology, in spite of Lankester"s declaration that this belonged to an older "Platonic" philosophy, and ought to be superseded by a term more consonant with the new philosophy of evolution.
[366] _Generelle Morphologie der Organismen. Allgemeine Grundzuge der organischen Formenwissenschaft, mechanisch begrundet durch die von Ch. Darwin reformierte Descendenztheorie_. Berlin, 1866. Reprinted in part as _Prinzipien der generellen Morphologie der Organismen_.
Berlin, 1906.
[367] He mentions as his predecessors in this field, Bronn, J. Muller, Burmeister, and G. Jager.
[368] In _Grundriss einer Allgemeinen Naturgeschichte der Radiolarien_, Berlin, 1887, and _Kunstformen der Natur_, Suppl. Heft, Leipzig.
[369] Haeckel had an intense admiration for Goethe"s morphological work. It is a curious coincidence that the work of Goethe, Oken and Haeckel was closely a.s.sociated with the town of Jena.
[370] But he himself would not admit this! See _Gen.
Morph._, ii., p. 11.
[371] _Fur Darwin_, 1864. Eng. trans, by Dallas as _Facts and Arguments for Darwin_, London, 1869.
[372] The bion is the physiological, as the morphon is the morphological, individual.
[373] See Vogt, _Embryologie des Salmones_, p. 259, 1842, and _supra_, p. 230.
[374] _An Essay on Cla.s.sification_, London, 1859.
[375] It was hinted at by Tiedemann. "It is clear that, proceeding from the earlier to the more recent strata, a gradation in fossil forms can be established from the simplest organised animals, the polyps, up to the most complex, the mammals, and that accordingly the animal kingdom as a whole has its developmental periods just like the single individual organism. The species and genera which have become extinct during the evolutionary process may be compared with the organs which disappear during the development of the individual animal" (p. 73, 1808).
[376] _The History of Creation_, vol. i., p. 310, 1876.
Translation of the _Naturliche Schopfungsgeschichte_, 1868.
[377] _Cf._ a parallel pa.s.sage from Serres, _supra_, p.
82.
[378] _Jenaische Zeitschrift_, ix., pp. 402-508, 1875.
[379] _Loc. cit._, ix., p. 409.
[380] _Untersuchungen zur vergl. Anatomie d.
Wirbelthiere_, Leipzig, i., 1864; ii., 1865; and iii., 1872.
[381] "U. d. Biologie in Jena wahrend des 19 Jahrhunderts," _Jenaische Zeitschrift_, x.x.xix., pp.
713-26, 1905.
[382] _Grundriss der vergl. Anatomie_, 1874, 2nd ed., 1878. Trans. by F. Jeffrey Bell, revised by E. Ray Lankester, as _Elements of Comparative Anatomy_, London, 1878.
[383] "This theory (evolution) shows that what was formerly called "structural plan" or "type" is the sum of the dispositions (_Einrichtungen_) of the animal organisation which are perpetuated by heredity, while it explains the modifications of these dispositions as adaptive states. Heredity and adaptation are thus the two important factors through which both the unity and the variety of organisation can be understood"
(_Grundzuge_, p. 19).
[384] _History of Creation_, i., pp. 241-2.
[385] "On the use of the term h.o.m.ology in Modern Zoology, and the distinction between h.o.m.ogenetic and h.o.m.oplastic agreements," _Ann. Mag. Nat. Hist._ (4), vi., pp. 35-43, 1870.
CHAPTER XV
EARLY THEORIES ON THE ORIGIN OF VERTEBRATES
Haeckel and Gegenbaur set the fashion for phylogenetic speculation, and up to the middle "eighties, when the voice of the sceptics began to make itself heard, the chief concern of the younger morphologists was the construction of genealogical trees. The period from about 1865 to 1885 might well be called the second speculative or transcendental period of morphology, differing only from the first period of transcendentalism by the greater bulk of its positive achievement. It must be remembered that the later workers (at least towards the end of this period) had immense advantages over their predecessors in the matter of equipment and technique; they possessed well-fitted laboratories in the university towns and by the sea; they had at their command perfected microscopes and microtomes; while the whole new technique of microscopical anatomy with its endless variety of stains and reagents made it possible for the tyro to confirm in a day what von Baer and Muller had taken weeks of painful endeavour to discover.[386] But the democratisation of morphology which followed upon the facilitation of its means of research left an evil heritage of detailed and unintelligent work to counterbalance the very great and real advances which technical improvements alone rendered possible.
This period of rapid development, which set in soon after the coming of evolution and multiplied the concrete facts of morphology an hundredfold, may for our present purpose be conveniently divided into two somewhat overlapping periods, of which the second may be said to begin with the enunciation by Haeckel of his Gastraea theory. Within the first period fall the evolutionary speculations a.s.sociated with the names of Kowalevsky, Dohrn, Semper, and others; the characteristic of the second period is the preponderating influence exercised upon phylogenetic speculations by the germ-layer doctrine in its two main evolutionary developments, the Gastraea and Coelom theories.
In the first period we might again distinguish two main tendencies, according as speculations were based mainly upon anatomical or mainly upon embryological considerations, and it so happens that these two tendencies are very well ill.u.s.trated by the various theories as to the origin of Vertebrates which began to appear towards the "seventies. We shall accordingly, in this chapter, consider very briefly the history of the earlier views on the phylogeny of the vertebrate stock.
In the early days, before the other claimants to the dignity of ancestral form to the Vertebrates--_Balanoglossus_, Nemertines and the rest--had put in an appearance, there were two main views on the subject, one upheld by Haeckel, Kowalevsky and others, to the effect that the proximate ancestor of Vertebrates was a form somewhat resembling the ascidian tadpole, the other supported princ.i.p.ally by Dohrn and Semper that Vertebrates and Arthropods traced their descent to a common segmented annelid or pro-annelid ancestor. The former view is historically prior, and arose directly out of the brilliant embryological investigations of A. Kowalevsky, who proved himself to be a worthy successor of the great comparative embryologist Rathke. His work was indeed a true continuation of Rathke"s. It was not directly inspired by evolution, though it supplied much useful confirmation of the theory--you may read Kowalevsky"s earlier memoirs and not realise that they were written several years after the publication of the _Origin of Species_.
His first paper of evolutionary importance was a note in Russian on the development of Amphioxus, published in 1865. This subject was followed up in two papers which appeared in 1867[387] and 1877.[388] In his papers on Amphioxus Kowalevsky made out the main features in the development of this primitive form, and showed that the chief organs were formed in essentially the same way as in Vertebrates; he described the formation of the archenteron by inv.a.g.i.n.ation, the appearance of the medullary folds, which coalesced to form the neural ca.n.a.l, the formation of the notochord and of the gill-slits. At first he made the mistake of supposing that the body-cavity arose from the segmentation-cavity, but in his later paper he rightly surmised that it was formed from the cavities of the "primitive vertebrae," or mesodermal segments. The origin of the notochord from the endoderm was also not made out by Kowalevsky in his paper of 1867.
Although many important details remained to be discovered by later investigators,[389] Kowalevsky"s work at once made the development of Amphioxus the key to vertebrate embryology, the typical ontogeny with which all others could be compared.
Meanwhile, in 1866 and 1871, Kowalevsky had communicated memoirs of even greater interest,[390] in which he showed that the simple Ascidians developed in an extraordinarily similar way to Amphioxus and hence to Vertebrates in general. His proof that Ascidians also develop on the vertebrate type aroused great interest at the time, and was naturally acclaimed by the evolutionists as a striking piece of evidence in favour of their doctrine. The systematic position of the Ascidians was at that time quite uncertain; they were grouped, as a rule, with the Mollusca, and certainly no one suspected that their well-known tailed larvae, first seen by Savigny, showed any but the most superficial a.n.a.logy with the tadpoles of Amphibia. Kowalevsky"s papers put a different complexion on the matter. In the first of them he showed how the nervous system of the simple Ascidian developed from ectodermal folds just as it did in Amphioxus and Vertebrates, how gill-slits were formed in the walls of the pharynx, and how there existed in the ascidian larva a structure which in position and mode of development was the strict h.o.m.ologue of the vertebrate notochord. In his second paper he entered into much more detail, and published some excellent figures, often reproduced since (see Fig. 13), but the proof of the affinity between Vertebrates and Ascidians was in all essentials complete in his paper of 1866.
[Ill.u.s.tration: FIG. 13.--Development of the Ascidian Larva. (After Kowalevsky.)]
Kowalevsky"s results were accepted by Haeckel, Gegenbaur, Darwin,[391]
and many others as conclusive evidence of the origin of Vertebrates from a form resembling the ascidian tadpole; they were extended and amplified by Kupffer[392] in 1870, later by van Beneden and Julin[393]
and numerous other workers; they were adversely criticised by Metschnikoff[394] and von Baer,[395] as well as by H. de Lacaze-Duthiers and A. Giard.[396] Lacaze-Duthiers and von Baer both held fast to the old view that Ascidians were directly comparable with Lamellibranch molluscs; they denied the h.o.m.ology of the ascidian nervous system with that of Vertebrates, von Baer being at great pains to show that the ascidian nerve-centre was really ventral in position.
He pointed out also that the "notochord" was confined to the tail of the ascidian larva. Giard"s att.i.tude was by no means so uncompromising, and the criticisms he pa.s.sed on the Kowalevsky theory are both subtle and instructive. He admits that there exists a real h.o.m.ology between, for instance, the notochord of Vertebrates and that of Ascidians. "But," he adds, "it is too often forgotten that h.o.m.ology does not necessarily mean an immediate common origin or close relationship. There exist, doubtless, h.o.m.ologies of great atavistic importance--I consider as such, for example, the formation of the cavity of Rusconi [the archenteron] in Ascidians and lower Vertebrates. But there are also adaptive and purely a.n.a.logical h.o.m.ologies, such as the interdigital palmation of aquatic birds, amphibians and mammals. These are not purely a.n.a.logous organs, for they can be superposed one on another, which is not the case with simply a.n.a.logous structures (the bat"s wing, for example, cannot be superposed on the bird"s wing); they are h.o.m.ologous formations, resulting from the adaptation of the same fundamental organs to identical functions. Such is, in my opinion, the nature of the h.o.m.ology existing between the tail of the ascidian tadpole and that of Amphioxus or of young amphibians. The ascidian larva, having no cilia and being necessarily motile, requires for the insertion of its muscles or contractile organs ... a central flexible axis, a true chorda dorsalis a.n.a.logous to that of Vertebrates" (pp. 278-9). This point of view is strengthened by the fact that in _Molgula_, studied by Lacaze-Duthiers, the embryo is practically stationary, and forms no notochord, nor ever develops sense-organs in the cerebral vesicle.
Giard"s general conclusion is that "the true h.o.m.ology with Vertebrates ceases after the formation of the cavity of Rusconi and the medullary groove: the h.o.m.ologies established by Kowalevsky for the notochord and the relations of the digestive tube and nervous systems are not atavistic, but adaptive, h.o.m.ologies" (p. 282). There is accordingly no close genetic relationship between Ascidians and Vertebrates.
Giard"s criticisms did not avail to check the vogue of the new theory, which soon became an accepted article of faith in most morphological circles.[397] The fall of the Ascidians from their larval high estate provided the text for many a Darwinian sermon.
Some years after the genetic relationship of Ascidians and Vertebrates had been established, a rival theory of the origin of Vertebrates made its appearance--a theory which was practically a rehabilitation in a somewhat altered form of the old Geoffroyan conception that Vertebrates are Arthropods walking on their backs. This was the so-called Annelid theory of Dohrn and Semper. Both Dohrn and Semper started out from the fact that Annelids and Vertebrates are alike segmented animals, and it was an essential part of their theory that this resemblance was due to descent from a common segmented ancestor. Both laid great stress on the fact that the main organs in Vertebrates are arranged in the same way as in an Annelid lying on its back, the nervous system being uppermost, the alimentary system coming next, and below this the vascular.
Dohrn"s earlier views are contained in the fascinating little book published in 1875, which bears the t.i.tle _Der Ursprung der Wirbelthiere und das Princip des Functionswechsel_ (Leipzig). He followed this up by a long series of studies on vertebrate anatomy and embryology,[398] in which he modified his views in certain details. We shall confine our attention to the first sketch of his theory.
If the Vertebrate is conceived to have evolved from a primitive Annelid which took to creeping or swimming ventral surface uppermost, a difficulty at once arises with regard to the relative positions of the "brain" and the mouth. In Vertebrates the brain, like the rest of the nervous system, is dorsal to the mouth and the alimentary ca.n.a.l; in an inverted Annelid, however, the brain is ventral to the mouth and is connected with the dorsal nerve cord by commissures pa.s.sing round the oesophagus. It would seem, therefore, that the primitive Vertebrate must have acquired either a new brain or a new mouth. Dohrn took the latter view. He supposed that the original mouth of the primitive ancestor lay between the _crura cerebelli_ in the _fossa rhomboidea_, and that in Vertebrates this mouth has been replaced functionally by a new ventrally placed mouth, formed by the medial coalescence of a pair of gill-slits.[399] Probably the two mouths at one period co-existed, and the older one was ousted by the growing functional importance of the newer mouth.
The gill-slits were considered by Dohrn to be derived from the segmental organs of Annelids, which were present originally in every segment of the primitive ancestor. The gills were at first external, like the gills of many Chaetopods at the present day. For their support cartilaginous gill-arches naturally arose in the body-wall, and the superficial musculature became attached to these bars. "There existed in all the segments of the Annelid-ancestors of Vertebrates gills with cartilaginous skeleton and gill-arches in the body wall. Each gill had its veins and arteries, each had its branch of the ventral nerve-cord, and between each successive pair of gills a segmental organ opened to the exterior" (p. 14, 1875). The paired fins and limbs of the Vertebrate arose by the functional transformation of two pairs of these gills. The anterior gills became the definitive internal gills of the Vertebrate, for they gradually shifted into the mouths of the anterior segmental organs, which had already acquired an opening into the pharynx and had been transformed into true gill-slits. The posterior gills degenerated and disappeared, but their arches remained as ribs. Gill-arches and ribs were accordingly h.o.m.ologous structures and formed a _parietal_ skeleton.
The vertebrate a.n.u.s, like the mouth, was probably secondary and formed from a pair of gill-slits, the post-a.n.a.l gut of vertebrate embryos hinting that the original a.n.u.s was terminal as in Annelids. The unpaired fins of fish were originally paired and possibly arose from the coalescence of rows of parapodia. Dohrn a.s.sumed also that the primitive Annelid ancestor must have possessed a notochord to give support in swimming.
If Vertebrates arose from primitive Annelid ancestors, how account for Amphioxus and the Ascidians, which seem to be the most primitive living Vertebrates and yet show no particular annelidan affinities? Dohrn tries to answer this awkward question by showing that these forms are not primitive but degenerate. He points out first that Cyclostomes are degenerate fish, half specialised and half degraded in adaptation to a parasitic mode of life. He thinks that if an _Ammocoetes_ were to become s.e.xually mature and degenerate still further, forms would result which would resemble Amphioxus, and ultimately, if the process of degeneration went far enough, larval Ascidians. Amphioxus therefore might well be considered an extremely simplified and degenerate Cyclostome, and the ascidian larva the last term of this degeneration-series. Both Amphioxus and the Ascidians would accordingly be descended from fish, instead of fish being evolved from them.
Dohrn conceived that the transformation of the Annelid into the Vertebrate took place mainly by reason of an important transforming principle, which he calls the principle of function-change. Each organ, Dohrn thinks, has besides its princ.i.p.al function a number of subsidiary functions which only await an opportunity to become active. "The transformation of an organ takes place by reason of the succession of the functions which one and the same organ possesses. Each function is a resultant of several components, of which one is the princ.i.p.al or primary function, while the others are the subsidiary or secondary functions. The weakening of the princ.i.p.al function and the strengthening of a subsidiary function alters the total function; the subsidiary function gradually becomes the chief function, the total function becomes quite different, and the consequence of the whole process is the transformation of the organ" (p. 60). Examples of function-change are not difficult to find. Thus the stomach in most Vertebrates performs both a chemical and a mechanical function, but in some forms a part of it specialises in the mechanical side of the work and becomes a gizzard, while the remaining part confines its energies to the secretion of the gastric juice. So, too, it is through function-change that certain of the ambulatory appendages of Arthropods have become transformed into jaws--their function as graspers of food has gradually prevailed over their main function as walking limbs. In the evolution of Vertebrates from Annelids the principle came into action in many connections--in the formation of a new mouth from gill-slits, in the transformation of gills into fins and limbs, of segmental organs into gill-slits, and so on.
Dohrn tells us that the principle of function-change was suggested to him by Mivart"s _Genesis of Species_ (1870), and he points out how it enables a partial reply to be made to the dangerous objection raised against the theory of natural selection that the first beginnings of new organs are necessarily useless in the struggle for existence.
We may note in pa.s.sing that a somewhat similar idea was later applied by Kleinenberg to the explanation of some of the ancestral features of development. He pointed out in his cla.s.sical memoir on the embryology of the Annelid _Lopadorhynchus_[400] that many embryonic organs seem to be formed for the sole purpose of providing the necessary stimulus for the development of the definitive organs. Thus the notochord is the necessary forerunner of the vertebral column, cartilage the precursor of bone. "From this point of view," he writes, "many rudimentary organs appear in a different light. Their obstinate reappearance throughout long phylogenetic series would be hard to understand were they really no more than reminiscences of bygone and forgotten stages. Their significance in the processes of individual development may in truth be far greater than is generally recognised. When in the course of the phylogeny they have played their part as intermediary organs (_Vermittelungsorgane_) they a.s.sume the same function in the ontogeny.
Through the stimulus or by the aid of these organs, now become rudimentary, the permanent parts of the embryo appear and are guided in their development; when these have attained a certain degree of independence, the intermediary organ, having played its part, may be placed upon the retired list."[401]
Dohrn was well aware of the functional, or as he calls it, the physiological, orientation of his principle, and he rightly regarded this as one of its chief merits. He held that morphology became too abstract and one-sided if it disregarded physiology completely; he saw clearly that the evolution of function was quite as important a problem as the evolution of form, and that neither could be solved in isolation from the other. "The concept of function-change is purely physiological;" he writes, "it contains the elements out of which perhaps a history of the evolution of function may gradually arise, and for this very reason it will be of great utility in morphology, for the evolutionary history of structure is only the concrete projection of the content and course of the evolution of function, and cannot be comprehended apart from it" (p. 70).[402]