A Historical Perspective on Vavilov’s Law of Homologous Series

Abstract About one hundred years ago, the law of homologous series, a sweeping generalization proposed by N. I. Vavilov at a plant breeders’ congress, caused a sensation, so much so that the audience thought they had witnessed the unveiling of something as revolutionary as the periodic table of Mendeleev. Vavilov postulated that the enormous diversity observable in plant forms masked an underlying similarity, one revealed in parallel variations among closely or distantly related plants. In the present article, we consider the law of homologous series from a historical viewpoint and conclude that its unveiling deserves to be recognized as a foundational event in evolutionary developmental biology. Similar content being viewed by others Data Availability Not applicable. Notes The periodic table, formally proposed by D. I. Mendeleev (1834–1916) in 1869, is a two-dimensional arrangement of the chemical elements. As was to emerge subsequently, the arrangement itself brings out the reason for the recurring similarities displayed by the elements; besides, it has been a powerful predictive tool. Vavilov had the same hopes for his law. It has been noted that the structure of the periodic table resembles that of the two-dimensional array of linguistic vocalisations constructed by the Sanskrit grammarian Pāṇini (fourth century BCE, conjectured), and that the coincidence is not accidental (Ghosh and Kiparsky 2019). Alphabets in many Indian languages continue to be written as 2-D arrays, and their vocalisations are taught accordingly. Crucifers are leafy green vegetables (e.g., cauliflower, cabbage). Papavers are herbaceous plants (e.g., poppy). Solanaceae are nightshades (e.g., tomato, potato, aubergine). Cucurbitaceae belong to the gourd family (e.g., squash, pumpkin, melon). Chenopodiaceae belong to the subfamily of Amaranthaceae (e.g., beet). Caryophyllaceae are herbaceous plants of the carnation family (e.g., Dianthus, Silene). Homologous variations in clades of bdelloid rotifers are of particular interest for the basis of the Law, because these organisms are believed to have evolved and diversified in spite of being asexual (see Fontaneto et al. 2007). Their existence suggests a role for developmental noise and multiple stable states in the development of polymorphisms (see Vogt 2015). Eimer (1843–1898) had found striking resemblances in the forms of snail shells belonging to entirely different groups, and named the phenomenon ”Unabhängige Entwicklungsgleichheit/ Homöogenesis“. Based on her studies of butterfly wing patterns, his student Maria Grāfin von Linden (1869–1936) further stressed its importance. As Vavilov mentions later, from her studies on snail shells and butterfly wings, she found parallel patterns across species “without any relation to their affinity and biological surroundings.” (Vavilov 1922, p. 86). Like Eimer, she too got into disputes with selectionists (see Linden and Minot 1897). (The article is made up of two notes with opposite interpretations.) Lotsy was a strong advocate of hybridization, meaning crosses between species or (in Vavilov’s language) Linneons, as an equivalent to mutation, because both could give rise to heritable variations and new combinations of characteristics. For him, hybridization was the main route for the origin and evolution of species. Later, Vavilov points out that his Law holds good not just for the existing diversity of forms, but also for new forms that appear by hybridization or mutation. Vavilov sees mimicry as an illustration of the law of homologous variation. Punnett was not arguing against natural selection, only saying that sometimes a single mutation could lead to a major phenotypic change, on which selection would act. He asked, if mimicry was due to the action of many genes of small effect, why could they not be separated by recombination? The answer only became apparent much later (see Charlesworth 2016). They include T. H. Morgan and his associates (Drosophila), E. B. Babcock (the walnut, Juglans), E. Baur, who had noticed them in several related species of plants and animals, P. A. Saccardo (mushrooms and fungi), and Hugo de Vries (for noticing the general feature). St. G. J. Mivart (1827–1900) initially supported, and later strongly opposed, natural selection. According to Vavilov, it was Mivart who coined the expression “The law of homologous variation”. A reading of Mivart’s book (Mivart 1871) shows that he distinguishes between homologous and analogous traits in the manner one would today, and elsewhere uses “homologous” in the broader sense of similar. For example, “…parts are often homologous which have no direct genetic relationship…which ‘Natural Selection’ pure and simple, seems unable to explain” (Mivart 1871, p. 106). Later, while alluding to the feathers which are sometimes seen on the toes of fowls and pigeons, his language indicates that he had taken for granted that there was a law of homologous variation. Without explicitly enunciating the law, he says “…the law of homologous variation has led to the development of feathers on the legs, in a position corresponding with those on the wing, namely, on the outside of the tarsi and toes. I am strengthened in this belief [i.e., belief that there is such a law] by the following curious case of correlation, which for a long time seemed to me utterly inexplicable, — namely, that in pigeons of any breed, if the legs are feathered, the two outer toes are partially connected by skin. […] the very same digits which in the wing are completely united by skin become partially united by skin in the feet…” (p. 204). But here he seems to be alluding to serial homology, not homology between parts in organisms belonging to different species. In order to settle the point, he would have had to trace character evolution by carrying out phylogenetic analyses. The allusion to Quetelet may refer to his pointing out that regular, predictable phenomena at the level of aggregates are not inconsistent with significant variation among individuals; he showed that the mathematical “law of errors”—the Gaussian distribution—is applicable to physical and social variables too (see Jahoda 2015). An Internet search shows that undergraduate curricula at agriculture and forestry universities in India mention the Law, and it is also part of the syllabus for civil services examinations in agriculture and forestry. Badyaev’s opinion, expressed in a book review, contains much that is relevant to how Vavilov’s law has been viewed, and is exactly the opposite (Badyaev 2010), Vavilov always refers to comparisons between varieties. The word is used in the botanical sense and stands for a genotype or strain of a species. Whether two varieties of the same species are different is judged, among other ways, by carrying out crosses. He does not consider variation between individuals of the same genotype. Synteny means a correspondence between parts of genomes, chromosomes or chromosomal regions of different species. One sign of synteny is the presence of the “same” genes in the same sequence (Hammer and Schubert 1994). If there is competition, there will be no such association. The most likely outcome is the elimination of the least functionally useful module, unless it performs (other) irreplaceable regulatory functions. As genomic projects have shown, the regulatory component of the genome is often redundant, implying that there is always a nonzero probability of competition between modules. The interpretation is based on the hypothesis of slowly coevolving memberships of amino acid sets, which, according to them, provides a molecular-level explanation of the homology-versus-phylogenetic distance aspect of Vavilov’s law (Rogozin et al. 2008). An earlier study using the same method concluded that arthropods belonged to the same clade as deuterostomes, separate from the ecdysozoans. That inference has been disputed; see Telford et al. 2008). Vavilov offered an explanation later (Vavilov 1931). In this context, considering that Vavilov emphasizes the importance of viewing plant and animal forms as geometrical entities, it is noteworthy that he does not mention D’Arcy Thompson, who did not think natural selection explained the evolution of biological form (his book “On Growth and Form” had appeared in 1917). Vavilov always proclaimed that he was a Darwinist, and it may have been too much to identify himself with a prominent anti-Darwinist (in a sense). It has been suggested that the convergent evolution of forms between the prokaryote Myxobacteria and eukaryote Dictyostelid amoebae might have a physicochemical basis (del Angel et al. 2020). For a comparison with contemporary views of plant development see Dornelas and Dornelas (2005). A similar caveat applies in the case of analogy. For example, the skin is not considered to be an analogue of the lungs and the mouth, although in amphibians the skin contributes to respiration and to the absorption of water. It is quite obvious that differences in mechanisms of respiration and water absorption require different vectors of selection for amphibian skin, lungs, and mouth, although some functions may be identical. Skin, lungs, and mouth are functionally overlapping, though not analogous, organs. Similar to the theory underlying Vavilov’s law, Alexei A. Zavarzin (1886–1945) developed a theory of parallel structural variations in animals, in particular with regard to histology. His ideas seem like an anticipation of the Human and other Cell Atlas enterprises (see Zavarzin 1925). Dmitri N. Sobolev (1872–1949) too thought there were parallelisms in forms on account of underlying physical principles of development. He proposed that the broad features of evolutionary and ecological change could be explained on the basis of biogeochemical feedbacks, believed in orthogenesis and nomogenesis, and argued that environmental factors could lead to catastrophic changes in ecosystems (see Popov 2008; Nanjundiah et al. 2022). Just what “Modern Synthesis” stands for, deserves debating. Should it go beyond the fusion of natural selection with Mendelian genetics, should it include stochastic effects based on small population size (drift), should it include macromutation, etc.? Not everyone agrees on the answers. See Rao and Nanjundiah (2017) for a discussion, centred on the writings of J B S Haldane, who is commonly viewed as one of the architects of the Modern Synthesis. Haldane’s view of evolutionary mechanisms went well beyond natural selection and drift. The 1987 publication is an English-language translation of Vavilov’s writings on centers of origin. The idea was first proposed in two Russian-language publications of 1924 and 1926 (also see Nanjundiah et al. 2022). A secondary center could form where a species had spread initially from the origin, and the plants there showed recessive traits, according to Vavilov. And yet it could display parallel series of traits between strains of the same species. Therefore, the Law applies among strains of a species, between species, and between larger taxonomic units. We repeat that this was supposedly in Darwin’s mind. The extended quote is well known and comes towards the end of the concluding chapter: “[.] these elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have all been produced by laws acting around us. These laws, taken in the largest sense, being Growth with Reproduction; Inheritance which is almost implied by reproduction; Variability from the indirect and direct action of the external conditions of life, and from use and disuse; a Ratio of Increase so high as to lead to a Struggle for Life, and as a consequence to Natural Selection, entailing Divergence of Character and the Extinction of less-improved forms.”; (Darwin 1859, p.490). Also see Bradley 2022). See definition on Wikipedia: https://en.wikipedia.org/wiki/Orthogenesis; also discussed in Levit and Olsson (2006). Nomogenesis holds that evolution is a lawful process, not one that occurs via chance and natural selection. Berg’s book on nomogenesis came out in Russian in 1922 (also see Melkikh 2026). In a not uncritical introduction to the 1926 English translation (Berg 1926), D’Arcy Thompson writes “[s]ome forty years ago, speaking to an Oxford meeting of the British Association on some of the ‘difficulties of Darwinism,’ I used the phrase ‘laws of growth’ in very much the same sense (though with a stronger leaning to the physical side) as Dr. Berg means by his word ‘Nomogenesis.’” (Berg 1926, p. VII). Haldane (1932a, p. 12) says nomogenesis is based on “[v]ariation due to internal causes, but not at random”, and adds that Berg has written “by far the best anti-Darwinian book of the century”. The word was coined by Haeckel, but its present meaning owes more to de Beer, Garstang and others. A Wikipedia article on heterochrony is informative on this and other points: https://en.wikipedia.org/wiki/Heterochrony. Also see Keyte and Smith (2014). See Bonner (2004) for an example in the case of Cope’s rule. The internalist and externalist attitudes are discussed in Linde-Medina (2010) and Linde-Medina (2011). The second article also argues that there is no dichotomy between the two, indeed that there is only one meaningful perspective, namely natural selection. As to whether the internalist perspective is not in accord with Modern Synthesis, it all depends on one’s view of what the Modern Synthesis entails (see footnote 27). Berg (1926) provides many more instances of similar parallelisms. D’Arcy Thompson (1917, 1942) had earlier argued forcefully for environmental forces and the mechanical principles that operated in organismal bodies, not natural selection, as preeminent for the evolution of form. One could say that not just the basic form, the signal, recurs across species, but variations from it, the noise, also recur. Interesting links suggest themselves to the theme of phenotypic plasticity. (see Nanjundiah 2020). 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Additional information Publisher’s Note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Rights and permissions Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law. About this article Cite this article Nanjundiah, V., Geeta, R. & Suslov, V.V. A Historical Perspective on Vavilov’s Law of Homologous Series. Biol Theory (2026). https://doi.org/10.1007/s13752-026-00551-w Received: Accepted: Published: Version of record: DOI: https://doi.org/10.1007/s13752-026-00551-w