Supernumerary Spikelet Wheat Forms as Models for Studying Genetic Regulation of Inflorescence Development

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Supernumerary Spikelet Wheat Forms as Models for Studying Genetic Regulation of Inflorescence Development O. B. Dobrovolskayaa, b, * a

Federal Research Center Institute of Cytology and Genetics, Siberian Branch, Russian Academy of Sciences, Novosibirsk, 630090 Russia b Federal Research Center Vavilov All-Russian Institute of Plant Genetic Resources, St. Petersburg, 190000 Russia *e-mail: [email protected] Received May 9, 2020; revised June 4, 2020; accepted June 16, 2020

Abstract—Almost all wheat species have an unbranched spike. Tetraploid rivet wheat Triticum turgidum L., branched forms of which are widespread and have been known for about 2000 years, is an exception. As for other wheat species, supernumerary spikelet forms are rare, and supernumerary spikelet/branched spike belongs to nonstandard morphotypes. As one of the examples illustrating the law of homologous series in variation, N.I. Vavilov presented the trait “spike branching” peculiar “not only to many wheat and rye species but also to many other genera with spike inflorescence or panicle.” The studies of genetic factors underlying the formation of “supernumerary spikelet/spike branching” trait and the study of peculiarities of the development of inflorescences of nonstandard branched wheat forms made it possible to demonstrate a genetic nature of hereditary variation of this trait. At the same time, the group of supernumerary spikelet/branchedlines is heterogeneous, and different genetic mechanisms can underlie the formation of branched spike. This review presents a retrospective of scientific studies devoted to the creation of supernumerary spikelet wheat forms and to the study of genetics of “supernumerary spikelet/spike branching” trait and demonstrates the results of modern studies of genetic regulation of morphogenesis of cereal inflorescences using supernumerary spikelet lines as genetic models. Keywords: homologous series in variation, inflorescence, spike, spike branching, supernumerary spikelets, morphogenesis DOI: 10.1134/S1022795420110034

INTRODUCTION Studying a vast diversity of cultivated plants and their wild relatives, Nikolai Ivanovich Vavilov for the first time drew attention to genetically determined similarity and relationship of the traits within the species, as well as larger taxa (plant genera and families). Identification of patterns in manifestation of polymorphism and establishment of polymorphism classes by analogy with homological series in organic chemistry led to the discovery of the law of homologous series in variation. Nikolai Ivanovich formulated the law and for the first time reported it at the Third Congress of Breeders in Saratov in 1920. The law was repeatedly published: initially as a small report in the proceedings of the Third Congress of Breeders, then in 1922, the article was published in the Journal of Genetics, and it was published in expanded version in Theoretical Foundations of Breeding in 1935 [1]. In the century since the discovery of the law of homologous series in va