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. 2021 Nov 9;11:21995. doi: 10.1038/s41598-021-01483-0

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© The Author(s) 2021

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

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Schematic representation of the expression patterns of integument development genes, in three different species. (a) Arabidopsis thaliana (WUS by Grobeta-Hardt¹⁶; BELL1 by Robinson-Beers et al.⁵⁸; ANT by Elliot et al.¹⁵; KAN by Leon-Kloosterziel et al.⁴⁴; Eshed et al.⁴⁰; UCN by Enuguttii et al.⁵⁹). (b) Gnetum gnemon previously published (WUS by Nardmannn et al. 2009; Melbel1, GneANT, GnmoKANs and GnmoUCNs by Zumajo-Cardona and Ambrose¹⁹). (c) Ginkgo results presented here. (d) Illustration of telome theory, synangial hypothesis and neo-synangial hypothesis for the origin of the seed. Notably, the telome theory indicates the evolution of integuments from sterile structures while both the synangial and neo-synangial hypotheses indicate the evolution of integuments from fertile (sporangia) structures.