Abstract
The concept of chordates arose from the alliance between embryology and evolution in the second half of the nineteenth century, as a result of a theoretical elaboration on Kowalevsky’s discoveries about some fundamental similarities between the ontogeny of the lancelet, a putative primitive fish, and that of ascidians, then classified as molluscs. Carrying out his embryological studies in the light of Darwin’s theory and von Baer’s account of the germ layers, Kowalevsky was influenced by the German tradition of idealistic morphology that was concerned with transformations driven by laws of form, rather than with a gradual evolution occurring by means of variation, selection and adaptation. In agreement with this tradition, Kowalevsky interpreted the vertebrate-like structures of the ascidian larva according to von Kölliker’s model of heterogeneous generation. Then, he asserted the homology of the germ layers and their derivatives in different types of animals and suggested a common descent of annelids and vertebrates, in agreement with Saint-Hilaire’s hypothesis of the unity of composition of body plans, but in contrast with Haeckel’s idea of the Chordonia (chordates). In The Descent of Man Darwin quoted Kowalevsky’s discoveries, but accepted Haeckel’s interpretation of the ascidian embryology within the frame of a monophyletic tree of life that was produced by the fundamental biogenetic law. Joining embryology to evolution in the light of idealistic morphology, the biogenetic law turned out to be instrumental in bringing forth different evolutionary hypotheses: it was used by Haeckel and Darwin to link vertebrates to invertebrates by means of the concept of chordates, and by Kowalevsky to corroborate the annelid theory of the origin of vertebrates. Yet, there was still another interpretation of Kowalevsky’s discoveries. As an adherent to empiricism and to Cuvier’s theory of types, von Baer asserted that these discoveries did not prove convincingly a dorsal position of the nervous system in the ascidian tadpole larva; hence, they could not support a homology between different animal types suggesting a kinship between ascidians and vertebrates.
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Notes
French original: “Il y a eu littéralement, de 1870 à 1900, une débauche, parfois même un délire de phylogénie, dont la responsabilité remonte surtout à Haeckel”.
French original: “Le système nerveux est, au fond, tout l’animal; les autres systèmes ne sont là que pour l’entretenir et le servir”. Annales du Muséum d’Histoire Naturelle 19, 1812, p 76.
In: Heuss T (1991) Anton Dohrn. A life for science. Springer, Berlin, pp 179–180.
German original: “So wandelt eine solche unwissenschaftliche Vergleichung wie in einem Labyrinthe, in dem an den ersten Irrweg nur neue sich anreihen”.
French original: “Je n’aurais pas parlé ici de ces conceptions, si elles étaient restées dans le domaine de ces théories abstraites dont on nous a gratifiés à foison depuis un certain temps et qui trouveront leur fin comme la défunte philosophie de la nature. Mais on se heurte à chaque pas à ces divagations; et elles se mêlent, chez certains auteurs, tellement avec les faits observés, qu’il est souvent difficile de démêler les éléments de la mixture qu’on vous offre. Il y a en outre un danger sérieux…”. Sur un nouveau genre de Médusaire sessile, Lipkea ruspoliana (C.V.), 1887, p 37, quoted by de Quatrefages (1894), p 119
German original: “Die wichtigsten Wahreiten in den Naturwissenschaften sind wederallein durch Zergliederung der Begriffe der Philosophie, noch allein durch blösses Erfahren gefunden worden, sondern durch eine denkende Erfahrung welche das Wesentliche von dem Zenfälligen unterscheidet, und dadurch Grundsätze findet, aus welchen viele Erfahrungen abgeleitet werden. Dies ist mehr als blösses Erfahren, und wen Man will, eine philosophische Erfahrung”. Handbuch der Physiologie des Menschen für Vorlesungen. Vol 2, p 522.
Contrary to Kowalevsky’s description, only in echinoderms the blastopore becomes the anus, in Phoronis, in particular, it closes towards anterior and its residual opening gives rise to the mouth.
As reported by primary and secondary scientific literature (e.g. Willey 1893; Dawydoff 1928; Brien 1948), Kowalevsky described the primordia of the branchial atria of ascidians as if they took their origin from two ectodermal invaginations. However, according to Kowalevsky the atrial cavities arise at first as evaginating pouches of the anterior gut. In his paper on Perophora listeri (Kowalevsky 1870a, translated from Russian by A. Giard 1874) he said: “…on voit qu’il est possible de comparer entre elles, et même de considérer comme parties homologues, la cavité du corps des Échinodermes et la cavité cloacale des Ascidies. Chez les Ascidies, il y aurait donc, pendant toute la vie, deux sortes de cavités, dont l’une, provenant de la cavité de segmentation, répondrait à la fentre qu’on remarque chez la Sagitta entre le feuillet externe et le feuillet interne; l’autre, la cavité cloacale, serait l’homologue de la cavité du corps de la Sagitta, des Échinodermes et de certains autres animaux”. In a similar way, in his paper on Ciona (Ascidia) intestinalis (1871a) Kowalevsky asserted that the greatest part of the branchial atria was endodermal, while the two ectodermal invaginations gave rise to the rudiment of the cloacal siphon: “Es bleibt uns noch jetzt zu erklären, welchen Antheil die Einstülpungen an der Bildung der Kiemenspalten nehmen… Man beobachtet an diesem Stadium, dass der obere Theil des Vorderdarms sich an beiden Seiten als zwei Falten erhebt, welche bald so gross werden, dass sie einen Theil der Gehirnblase von den Seiten bedecken und ihr hinteres Ende (Fig. 34, Tafel XII) jederseits dicht an die durch Einstülpung entstandene Kloake stösst”.
German original: “Dagegen möchte ich anführen, dass, wenn wir z. B. die Wirbelthiere, als einen überhaupt hoch organisirten Typus, von einem Urvater ableiten, der zu den niedrig stehenden Typen der Thiere gehörte z. B. zu den Mollusken (vielleicht Tunicaten) oder Würmern (z. B. Sagitta oder ähnlichen) so wären doch die Keimblätter der zuerst entstandenen Wirbelthiere mit denjenigen der anderen Typen zu vergleichen, und wenn wir die Keimblätter des Amphioxus mit denjenigen der Würmer und Mollusken vergleichen, so müssen wir dies mit den Keimblättern auch der anderen Wirbelthiere machen”.
Kowalevsky ended this paper by the following sentence: “Aus allen diesen Gründen halte ich die Ansicht, dass die Organe der Thiere verschiedener Typen nicht homolog sein könnten, für nich haltbar”.
German original: “Wenn eine solche Annahme auch sehr befremdend erscheint, so stimmt sie doch mit der jetzt herrschenden Anschauung, dass embryonale Bildungen, embryonale Stadien wirklich existirende Formen darstellen”.
German original: “Wenn Claparède noch zweifelt, ob diese Fasern doch noch vielleicht als Nervenfasern anzusehen sind, so ist ihre Entwicklung aus dem mittleren Blatte ein so wichtiger Grund gegen ihre Nervennatur, dass ich dieselben keineswegs für Nervenfasern ansehen kann”. (Kowalevsky 1871a, p 123).
In Darwin’s words: “If this work had appeared before my essay had been written, I should probably never have completed it. Almost all the conclusions at which I arrived I find confirmed by this naturalist, whose knowledge on many points is much fuller than mine. Wherever I have added any fact or view from Prof. Haeckel’s writings, I give his authority in the text; other statements I leave as they originally stood in my manuscript, occasionally giving in the foot-notes references to his work, as a confirmation of the more doubtful or interesting points”.
In The life and letters of Charles Darwin, vol 3, p 180.
In The life and letters of Charles Darwin, vol. 3, p 198. The year of this letter is uncertain and is reported as (1875?).
This account of Michael Foster was included in an anonymous review article: The kinship of Ascidians and Vertebrates. Q J Microsc Sci 10:59–69, 1870. It seems quite possible that the author of the review was Edwin Ray Lankester, the editor of the journal.
Evidently, von Baer was following the chordate affair with close attention, since he said that the paper of Dönitz, first expected from July 1870, was likely to be issued in late 1871, being included at the end of the 1870 volume of the Archiv für Anatomie und Physiologie. An English translation of the paper of Dönitz was published in 1871 in a review article, Q J Microsc Sci 9:281–283.
German original: “Müssen wir nicht hierin ein allgemeines Gesetz der bildenden Natur vermuthen?”.
Unlike Kowalevsky, Kupffer (1870) noticed the unusual dorsal position of the mouth in the ascidian larva, but he considered this detail to be of minor importance as compared to the vertebrate-like organization of the body. To account for this peculiarity, he suggested that the development of the adhesive papillae could shift the mouth to the dorsal side.
German original: “Der Lehre von der Transmutation der Thierformen principiell nicht abgeneigt, sondern eher zugeneigt, verlange ich doch vollständigen Beweiss, bevor ich an eine Umwandlung des Wirbelthier-Typus in den der Mollusken glauben kann”.
German original: “Man darf nun wohl vermuthen, dass überhaupt der Stoff für die Bildung der Centraltheile des Nervensystems aus der äussern Schicht der ersten Anlage des Embryos genommen wird und durch Einfaltung die ihm gebührende Stelle erhält, und dass diesem allgemeinen Gesetze gemäss auch das Nervencentrum der Tunicaten durch Einfaltung aus der äussern Embryonal-Schicht entsteht”.
The paper of Kowalevsky Embryologische Studien an Würmern und Arthropoden was submitted to the Physico-Mathematical Class of the Academy of Sciences of St. Petersburg on 18 November 1869 and published in 1871. Although he dissented from his conclusion, von Baer gave his positive vote to Kowalevsky also on occasion of the first von Baer-Prize in 1867, when the award was made to Kowalevsky for his study of ascidian development.
Kowalevsky (1871a) replied to Metschnikoff on several points concerning, in particular, the fate of the blastopore and the germ layer derivation of notochord and nervous system, but he never discussed the problem of orientation.
German original: “Ich hatte dabei die vielen Dilettanten im Auge, die an vollständige Transmutationen glauben, und die geneigt sein werden, es für blosse Eitelkeit zu halten, wenn man in den Ascidien nicht die Vorfahren der Menschen erkennen will”.
In his treatise on embryology (1828) von Baer said that if it were a bird writing such a volume in the light of the theory of transmutation, he would note that adult humans resemble embryonic birds. Just as the latter, humans lack beaks, have anterior and posterior limbs which are similar to each other, and moreover, they show tiny hairs which correspond to the most primitive form of feather production. As a conclusion, birds are higher forms which evolved from humans.
“Of late the attempt to arrange genealogical trees involving hypothetical groups has come to be the subject of some ridicule, perhaps deserved. But since this is what modern morphological criticism in great measure aims at doing, it cannot be altogether profitless to follow this method to its logical conclusions. That the results of such criticism must be highly speculative, and often liable to grave error, is evident”. In: The ancestry of the Chordata. Q J Microsc Sci 26, p 535–536.
Dealing with the problem of the origin of the notochordal cells of the lancelet Thomas H. Morgan wrote: “There has been much discussion as to whether these cells are to be called ectoderm or endoderm, but this discussion has lost interest since we have come to pay more attention to the cell-lineage of the embryo than to phylogenetic questions based on imaginary two-layered ancestors of those stages”. In: Experimental embryology, 1927, p 322.
Giard (1874) held that the kinship between ascidians and vertebrates shown by their developmental similarities was not so close as many people believed. French original: “Mais cette parenté est moins directe qu’on ne l’a supposé, et les points communs que l’on retrouve chez l’adulte indiquent bien moins une filiation qu’une certaine capacité à produire des formations de même nature, mais non synchrones, capacité qui est évidemment l’héritage d’un ancêtre éloigné commun”.
According to Carazzi (1906) “the vertebrates are to be considered separately from the lancelet from the embryological as well as the anatomical point of view… Even if we agree to use the inclusive term chordates for indicating tunicates, lancelets and vertebrates, we must take into account that tunicates and cephalochordates share a number of characters, but they differ widely from the vertebrates. This conclusion implies that the phylum Vertebrata is completely separated from all phyla of invertebrates”.
This ironic comment is taken from the scientific blog Pharyngula, run by P. Z. Myers and hosted on ScienceBlogs.com.
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The enduring support of the libraries of the “Anton Dohrn” Zoological Station of Naples and the Museum of Natural History “G. Doria” of Genoa is acknowledged with thanks.
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Raineri, M. On some historical and theoretical foundations of the concept of chordates. Theory Biosci. 128, 53–73 (2009). https://doi.org/10.1007/s12064-009-0059-y
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DOI: https://doi.org/10.1007/s12064-009-0059-y