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. 2023 May 4;9(5):720–732. doi: 10.1038/s41477-023-01405-0

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© The Author(s) 2023

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 license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license 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 license, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 7 — a, Model for the elaboration of the grass ligule and the leaf in which WOX3 function in the rim organizes tissue outgrowth between juxtaposed domains of adaxial and abaxial identity. Auxin transport is necessary for both leaf and ligule initiation, and Cuc2 boundary gene expression is present in both contexts. b–d, Proposed homologies for the ligule in monocots and eudicots (Arabidopsis): grass ligule and sheath margin are homologous to the margins of the eudicot leaf base and stipules (that is, the ligule is a stipule) (b), ligule and sheath margins are homologous to the margins of the eudicot petiole (that is, the ligule is an extended sheath margin) (c) and the ligule lacks homology with any eudicot leaf feature (d). e, Confocal fluorescence micrographs of compiled longitudinal sections through ligules at varying stages of maturity (left to right, from leaf). Cells express a pTUBULIN:CFP-TUBULIN construct. The length of the sheath at each stage is indicated (n = 3). Arrowheads indicate the ligule cleft. f, The BSB of a mature leaf. Arrowheads indicate the continuity between the rim of the sheath and the ligule. B, blade; L, ligule; S, sheath.