Abstract
In 1835, the meaning of the cleavage furrows in the division of frog eggs was the cause of a heated argument between the Italian naturalist Mauro Rusconi and Karl Ernst von Baer. These furrows were first described by Prévost and Dumas (Ann Sci Nat 2:100–121, 129–149, 1824b) who did not realize they cut the egg into separate masses. Rusconi (Développement de la grenouille comune depuis le moment de sa naissance jusque a son état parfait, Giusti, Milano, 1826) hypothesized a connection between the furrows and a peculiar crystallization of the content of the egg which eventually produced elementary molecules as the building blocks of the embryo. von Baer (Arch Anat Phys Wiss Med 6:481–509, 1834) was the first to establish a link between the furrows and an active process of dichotomous division he considered to be the basis for all further development and differentiation. The present paper analyses the theoretical reasons behind these divergent interpretations and focuses attention on their implications for the development of the cell theory and the conceptions of life. Prévost, Dumas and Rusconi interpreted cleavage and the whole embryonic development in the light of eighteenth-century scientific theories and the French materialism of the early nineteenth century, which explained life in terms of ordered molecular movement. Starting from other premises partly rooted in German philosophy von Baer (1834) gave a totally different picture which anticipated the cell theory and modern embryology.
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Reprinted with permission from the Handschriftenabteilung der Universitätsbibliothek Giessen, Nachlaß von Baer, Verschiedene Zeichnungen Embryos. Tafeln und Zeichnungen, Bd. 22, Bl. 19 (colour figure online)
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Notes
Beetschen quoted the German translation of Rusconi’s second letter to Ernst Heinrich Weber which was first published in an Italian periodic (Rusconi 1835b). In the present paper, all five of Rusconi’s letters are quoted in their original Italian version.
Rusconi's biographers reported that he did not always consider his words (Biffi 1853; Cattaneo 1925). A more realistic picture of the man was given by Verga (1869, pp. 41–52) in his account of the dispute between Rusconi and Bartolomeo Panizza (1785–1867). During this controversy which began in 1839 and was settled in 1844 by a commission that decided in favour of Panizza, Rusconi behaved like a true villain both in public and in private. Just as in the case of von Baer, he was greatly disturbed to see that Panizza had entered what he considered to be his personal field of research, i.e. the pattern of the lymphatic vessels in amphibians. Besides, he misinterpreted one of Panizza's sentences as an accusation of plagiarism. Concerning the development of frog eggs, in his five letters to Weber Rusconi (1835a, b, c, 1838, 1839) made a full list of accusations against von Baer: his boastful attitude to pontificate about the work of other researchers, while he himself was devoid of any skill when it came to carrying out scientific research; mental blindness owing to preconceptions and excessive imagination; ridiculous self-congratulatory praise of trivial results; extremely boring diffuse writing; sloppiness and malicious falsification of the quoted literature; ignorant petty physiologist and logician; liar; and so on. With reference to Rusconi’s second letter to Weber (1835b), Gasco (1880, p. 9) commented: "Rusconi assails and shakes him [von Baer] in such a way that on several occasions the reader of that incandescent letter feels shivers running down his spine and is going to let out: poor Baer!". In his autobiography, von Baer (1866, p. 380) said only a few words about this aggressive attitude: "… Ruscomi [sic] had been very much angered by my remark that he as well as Prévost and Dumas had observed division only on the surface, hence their usage of the term ‘cleavage’, an anger which he vented in Müllers Archiv, 136, p. 205, etc., and which, when I made his personal acquaintance 10 years later, I found still present". Most probably von Baer visited Rusconi in Pavia while travelling from Genoa to Venice via Milan in Autumn 1845. To our knowledge, this is the only available account of this face-to-face meeting, as Rusconi’s unpublished writings have been almost entirely lost. Rusconi died in 1849, but he still attacked von Baer in his final posthumously published work on amphibian development (Rusconi 1854)..
"As a consequence, it is fully demonstrated that the tadpoles exist before fertilization, a very interesting truth that I should like to prove for the sake of clarity in the following way. The unfertilized eggs do not differ at all from the fertilized ones; but the fertilized eggs are nothing else than the tadpoles concentrated and shrunk on themselves. It follows that the same must be held as to the unfertilized eggs. Thus, the tadpoles predate fertilization and in order to develop they only need the fertilizing fluid of the male" (Spallanzani 1768, p. 54).
In the introduction to his book, Rusconi pointed out that he had finished this study in 1821, but delayed its publication because he was not satisfied with the plates and wanted to draw and engrave them again. Therefore, the paper of Prévost and Dumas (1824b) was published first.
von Baer cited the principles of Cuvier as an argument against the Darwinian theory of evolution. Among other things, he based his criticism of the kinship between ascidians and vertebrates (von Baer 1873) on Cuvier’s embranchements, i.e. the existence of four distinct composition plans depending, in turn, on different patterns of the nervous system (Raineri 2009). In a letter to Anton Dohrn of 23 June 1875 (Groeben 1993, pp. 79–80), von Baer contrasted the theory of natural selection with Cuvier’s principles of the conditions of existence and of the final causes. "I am convinced that every normal development of an organism … must be regulated to make existence on this earth possible. Should an organism therefore, according to my opinion, be constructed that has to seek for its food on terra firma, then linked to this is the fact that it acquires jointed feet, and, what is more, according to the laws of mechanics". A few lines below he added: "I cannot break away from my old view that nature reasons and strives toward goals, whether immanently or transcendentally". While transcendental finality (Zweckmässigkeit) is a philosophical concept, immanent finality (Zielstrebigkeit) can be investigated by science (Kolchinsky and Levit 2019). Zielstrebigkeit, in fact, manifests itself in living organisms inasmuch as they are constructed mechanistically in order to exist in a given environment. von Baer (1866, p. 449) reasserted his position in the following comment to his own treatise on embryology (von Baer 1828, 1837): "My entire presentation, in both the above-mentioned volumes, has been reproached for being too mechanical. I admit that I accepted this reproach as a compliment, for one stands more firmly on a board than on aether or the sky at dawn". These nebulous concepts were likely to be the speculations of Naturphilosophie.
According to Pander (1817a, b), the developing germinal disc consisted at first of two layers, membrana pituitosa or mucous layer closer to yolk, and membrana serosa or serous layer. At a later stage, a third membrana vasculosa, or vascular layer, appeared between the two previous ones. In this treatise on embryology von Baer (1828) called serous and mucous layers animal and vegetative layers and reported that each of them differentiated, respectively, into skin and muscle layer, and vascular and mucous layer. von Baer (1828, 1834) acknowledged Pander’s priority in the discovery of the germ layers but rejected Pander’s idea that these layers and the spinal cord formed by a process of coagulation (von Baer 1866, pp. 299–300).
von Baer used the term "type" as a synonym of "developmental scheme" (Brauckmann 2011). Rather than being referred to an ideal archetype, the geometrical appearance of the type of vertebrates according to von Baer depended on his attempts to illustrate the three-dimensional development of the earliest embryonic rudiments by two-dimensional schematic drawings. In order to elaborate a general theory of development based on universal patterning laws, von Baer also intended to identify the types of invertebrates and prove the homology of their germ layers with those of vertebrates. To this end, he carried out preliminary studies of the development of insects, crustaceans, molluscs and worms. He also showed the developing egg of the river crayfish (Astacus fluviatilis) in four drawings in the plate of his Epistola on the mammalian egg (von Baer 1827a). On p. 24 of the same book, however, von Baer said a few words about an unpleasant affair he described at length in his autobiography (von Baer 1866, pp. 339–341). While he was cooperating with Burdach in the compilation of a treatise on physiology (Burdach 1828), von Baer told him privately that the development in arthropods is similar, but dorsoventrally inverted as compared to that of vertebrates, inasmuch as their embryo consists at first of a ventral axis and two bilaterally symmetrical plates which are thicker on the ventral than on the dorsal side. Without informing von Baer, Burdach communicated this observation to Martin Heinrich Rahtke (1793–1860), who was investigating the development of the river crayfish. Rathke (1825) then published his own results as soon as possible placing emphasis on the initial development of the crayfish from a ventral line. To the opposite, von Baer delayed his own publication since he knew that Rathke was working on the same subject and supposed that they could prepare a cooperative paper under the supervision of Burdach. Soon, however, he learnt that Rathke had proceeded to publish his preliminary results. von Baer was greatly disturbed by this misconduct and expressed some resentment in a letter to Rathke on 26 August 1826 (Universitätsbibliothek Giessen, Handschriftenabteilung, Nachlass von Baer, Bd. 25). In the same letter, however, von Baer credited Rathke with an excellent work that confirmed independently his own results. Rathke replied to von Baer that he knew nothing about his research on arthropods as Burdach had represented the hypothesis of their dorsoventrally inverted development as his own discovery. von Baer was later reconciled with Rathke, but his relationship with Burdach soured. In a subsequent study, Rathke (1829) described the germ layers of the crayfish and other developmental analogies between arthropods and vertebrates in greater detail and cited von Baer's 1827a preliminary observations on this subject.
Rusconi gave an account of oogenesis in the green frog in his work on the development of a land salamander. This study started in 1839, but was interrupted during the dispute with Panizza and only resumed in 1843. Rusconi, however, died in 1849 before publishing his results. The revised edition of the 1839 manuscript, including six plates from Rusconi’s original drawings, was published in 1854 by Joseph Morganti.
In the same study, Rusconi gave the first account of the modified neurulation in fish. In his words, "in fish at first the dorsal rod is not divided into two halves which are apart from one another, as found in frogs, salamanders and many other animals. In fish the back forms as a single piece altogether" (Rusconi 1835c, p. 256). Actually, in fish the neural plate sinks inside forming the neural keel which rounds up into a solid cylindrical rod. The neural tube is formed by cavitation at a later stage. By contrast, von Baer (1837) reported that neurulation in fish was nearly identical to that of amphibians, except from the initial position of the neural folds which were positioned further apart from one another. Almost certainly, he misinterpreted the germ ring as neural folds.
Following the publication of Ecker’s embryological atlas (1851–1859), the yolk plug was usually called Ecker’s Dotterpfropf or bouchon de Ecker.
Rusconi (1854, p. 36) asserted that "the law that was established by Serres is correct, unfortunately this famous anatomist has exaggerated, as far as we can judge, its importance and application". According to Serres (1827), the spinal cord was formed by apposition of successive layers, while Rusconi (1826) reported an origin by fusion of lateral folds. Another disagreement between the two naturalists concerned the anatomy of the larval nervous system.
von Baer came to disapprove of the definition ‘Dorsal chord, Chorda dorsalis’ that he used in his treatise on embryology (von Baer 1828). "I have always considered this chord to be specific to the central trunk, and whatever is formed above it to belong to the dorsal side, whatever is under it, to the abdominal side. I therefore would have preferred the word 'Vertebral chord, Chorda vertebralis’" (von Baer 1866, p. 448).
As he wrote in the introduction to his 1826 book, Rusconi had developed a special microscope to overcome the difficulty of investigating opaque objects and wanted to make it available to other students, including Prévost and Dumas as a sign of his esteem.
Concerning the Graafian follicle, von Baer was led into error by his tendency towards generalization and by the principle that differences at a later stage can be traced back to original conformity. As he did not regard it as a newly created structure that was present only in mammals, he considered the Graafian follicle (the maternal egg) to be the equivalent of the chicken egg. Accordingly, he interpreted the mammalian egg (the foetal ovum) and the granular layer- cumulus ooforus (which consist of follicle cells) as the analogues, respectively, of the Purkinje vesicle and the stratum proligerum-cumulus (germinal layer) of the egg of chickens and other vertebrates (von Baer 1827a, 1866).
For instance, on p. 87 of the Dictionnaire des Sciences Naturelles, vol. 58 (1829) we can read that: "These bodies, which have been classified by naturalists within the infusoria with the name of cercariae and are commonly called animalcules spermatiques, according to MM. Prévost and Dumas should be the first rudiment of the nervous system". The same classification within the infusoria under the name of von Baer was reported by Burdach’s treatise on physiology (1828, § 84).
Following the triumph of Darwinian theory, the prestige of Lamarck has been underestimated. However, "far from being universally scorned, Jean Baptiste Lamarck became known as ‘the French Linnaeus’ during the 1820s. Speaking at Lamarck’s funeral in December 1829, Étienne Geoffroy Saint-Hilaire remarked that the last years of the old naturalist’s life had been brightened by the awareness of how much his work was appreciated in Europe, and especially in France" (Corsi 2009, p. 167).
Leading exponents of the theory of the fibre were Albrecht von Haller (1757), Erasmus Darwin (1794) and Xavier Bichat (1801). According to Darwin (1794, p. 20), for instance, "the spirit of animation is the immediate cause of the contraction of animal fibres, it resides in the brain and nerves, and is liable to general or partial diminution or accumulation".
Carlo Chiolini quoted Amours des salamandres aquatiques (Rusconi 1821) and the still unpublished results on the development of the green frog to confute preformation and the above-mentioned theory of the fibre. According to this theory life began at fertilization, as the sperm (the agent part) elicited in the fibre (the reagent part) "the vital principle which is inherent to the nervous substance". On the contrary, Chiolini (1825, p. 66) agreed with Rusconi "to consider generation to be a kind of crystallization".
On the subject of amphibian artificial cross-fertilization, Rusconi (1838, p. 348) reported that "with the help of the sun heat the generative humour of the toad elicited in the molecules of the material of the egg [of frog] the internal movement which represents the organic life of the egg itself". While it was "slow and regular" in normal conditions, this movement was "irregular and turbulent" in cross-fertilized eggs which underwent abortive development.
With his usual precision, von Baer (1834, p. 503) asserted that he could not specify the extent to which the female generative fluid cooperated in the process of division. In any case, "it is clear that as soon as it is reached by the [male] generative substance diluted with water, the yolk sphere reveals itself as endowed with autonomous life. Previously it had only a latent life, as autonomous life will be aroused by artificial or natural fertilization with the same degree of certainty regardless the egg has remained for a short or long time in the dilated oviduct".
"It appears that the [first] furrow elongates in such a way that the mass of yolk retreats on both sides in opposite directions, in the meanwhile as the walls which are generated by the growing furrow form transitory folds and sometimes a slight, but clearly visible tremble runs across the adjoining mass of yolk. From this observation it clearly follows that the mass of yolk is not furrowed, as to say, by an invisible tool, but it splits into parts by a vital act" (von Baer 1834, p. 486).
Fifty years later, Hertwig (1884) asserted that cells divide along their long axes and put this rule into relationship with the central position of the nucleus and the orientation of the mitotic spindle parallel to the longest axis of the protoplasmic mass.
"As regards the subdivision of the cells, we have already seen how a jutting out of the cell-membrane may be produced by its more vigorous growth in certain situations. But a jutting inwards into the cavity of the cell may also result from the very same process. Now, if we imagine this jutting inwards to take place in a circular form around a cell, as the consequence of a partial increase in the force of its growth, it may proceed to such an extent, that one cell may be separated into two, connected together only by a short peduncle, which may afterwards be absorbed. This would illustrate the most simple form of subdivision in a cell. In animal cells, however, which undergo subdivision, that is, the fibre-cells, the process is more complicated; firstly, because when an elongated cell subdivides, it splits into many fibres, and, secondly, because the cells are so very minute. The process, therefore, cannot for these reasons be accurately traced, and the following is all we can detect: a cell becomes elongated on two opposite sides into several fibres; from the angle, which the fibres on either side form with each other, a striated appearance gradually extends over the body of the cell; this formation of striae becomes more and more distinct, until the body of the cell splits entirely into fibres" (Schwann 1839, p. 218).
According to several authors (e.g. Roger 1963; Bowler 1971), preformation in the broad sense of the word includes at least two theories. One, which may be called pre-existence, assumes that every living organism was formed by God since the Creation. An extreme version of this idea is the theory of encasement (emboîtement), which means that all future living individuals were encased within the reproductive organs of either the mother or the father from the beginning. Ovists and animalculists thought that either the egg or the sperm contained a miniaturized foetus. As noted by von Baer (1834), by the 1830s the theory of pre-existence had already been discredited, while many naturalists (including von Baer himself) still subscribed to some form of preformation, or the hypothesis that the future organism was preformed in the egg in a more or less organized state as a result of natural agents without the need of a divine intervention.
"In science the ideas become connected to one another; when a system prevails, its roots spread over its whole domain and give rise to opinions which often survive the hypotheses which generated them. The pre-existence of germs had as a result a sharp separation between organized and inorganic bodies, as the latter were thought to be totally devoid of germs …. Sciences became subdivided into two classes, organic and inorganic …. This is the origin of the ideas about the growth of the organized bodies and the elementary forms of the organs, which was claimed to be the exact opposite of the growth of the inorganic bodies. The latter occurs by juxtaposition: the molecules of growth place themselves around the original individual according to certain laws. The former should occur by intussusception; the primitive tissue expands itself and increases in size" (Serres 1827, pp. 59, 60).
The terms inducer and induction still used by embryologists to deal with organizing centres and embryonic patterning were introduced by Hans Spemann (1938) by analogy with magnetic induction. Just as von Baer, Spemann adopted a mechanistic approach even when he was not a reductionist himself. At the end of his 1938 book, he wanted to express his conviction "that the suitable reaction of a germ fragment, endowed with the most diverse potencies, in an embryonic ‘field’, its behaviour in a definite ‘situation’, is not a common chemical reaction, but that these processes of development, like all vital processes, are comparable, in the way they are connected, to nothing we know in such a degree as to those vital processes of which we have the most intimate knowledge, viz., the psychical ones” (Spemann 1938, p. 372).
"The fundamental point of every thing escapes us; we have investigated all non-essential difficulties, and we have solved them, until we came to the crux of the question, and then, suddenly the truth eludes all our efforts, their only usefulness being to testify to our impotence" (Dumas 1827, p. 454).
According to von Baer in amphibians just as in birds this protein-containing jelly was used for nourishment, at the larval or embryonic stage, respectively. He wondered how Rusconi (1826) could deny that tadpoles fed on particles of the jelly coat. Rusconi (1854) replied vehemently that the jelly coat did not provide any nourishment for the embryo, but he misquoted von Baer who had ascribed this feeding behaviour to larvae, not embryos of frogs.
"A few hours after fertilization a transformation begins, insofar as the upper side of the egg becomes divided by furrows which take their origin from the middle of the germinal layer and elongate on the brown hemisphere, and soon later on the clear one. The first furrow forms according to a diameter of the germinal layer, soon another one intersects it at right angle and a third one forms subsequently according to the horizontal section of the egg. Thus, the germinal layer comes to be subdivided by this horizontal furrow into two halves, each consisting of four segments. Other furrows form parallel to the initial ones and develop gradually, and then, they also are cut in the middle by transverse furrows. As a result, after approximately twelve hours the whole egg looks like a sharkskin. At a later stage, these furrows gradually disappear, and after sixteen hours, the egg is smooth again. After the furrows have disappeared, a tiny bump which may be called primitive streak is seen to rise in the middle of the germinal layer. Approximately at the thirtieth hour (as development goes on fast) on each side of the primitive streak other wider bumps arise, the dorsal plates" (Burdach 1828, p. 223). At page 297 of the 1837 edition of Burdach’s treatise the same paragraph authored by von Baer states that: "A few hours after fertilization a transformation begins, insofar as the upper side of the egg becomes divided by furrows which take their origin from the middle of the germinal layer and elongate on the brown hemisphere, and soon later on the clear one. After the furrows have disappeared a tiny bump which may be called primitive streak is seen to rise in the middle of the germinal layer etc". The description of the cleavage stage has been cut away altogether, while that of the initial formation of the embryo is the same of the 1828 edition. In the meantime, however, von Baer (1834, 1837) had changed his views on these topics significantly. Seemingly, he did not collaborate with Burdach in modifying this paragraph. Probably this split was a result of von Baer’s permanent settlement in St. Petersburg from 1834. On the other hand, he decided to shift to Russia also owing to frictions he had with Burdach while living in Königsberg (von Baer 1866, see note 8).
This preliminary note was cited by Remak (1852, p. 52) only in a footnote: "As for it concerns my observations on cell division, since several years Carl Ernst von Bär (Fror. N. Notiz 1846. No. 839) has expressed the opinion that the germinal vesicle coincides with the nucleus, the nuclei of the embryonic cells are the result of its division, and all cells and nuclei increase in number by division. When this article was already in press I turned my attention to this communication of Bär through Mr. Johannes Müller". Conversely, long before the paper of Virchow (1852) von Baer (1847) cited some preliminary observations of Remak on the origin of tissues by continuous division.
At the beginning of his 1847 note, von Baer (1847, p. 231) declared: "I have just finished a research which, as I believe, will leave its mark", though in a footnote he specified more modestly: "at least as for it concerns the experiments of artificial fertilization on marine animals". In any case, he felt that that the discovery of the egg division "brought me into the innermost tabernacle of embryology" (von Baer 1866, p. 377).
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Acknowledgements
We would like to thank Dr. Edoardo Razzetti (Natural History Museum, Pavia) for providing us with Rusconi’s 1826 and 1854 books as well as biographical information. Dr. Alberto Vianelli and Dr. Paolo Musso (University of Insubria, Italy) gave us bibliographic material on the history of the cell theory. We also thank the referees for their valuable comments which helped us to improve the manuscript.
Funding
Estonian Ministry of Education, Grant Numbers: P170021SPTL and P180276SPTL.
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Raineri, M., Tammiksaar, E. Scientific traditions in conflict: the Rusconi–von Baer controversy on the embryology of frogs and the development of the cell theory. Theory Biosci. 140, 45–75 (2021). https://doi.org/10.1007/s12064-020-00325-3
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DOI: https://doi.org/10.1007/s12064-020-00325-3