Reticulate leaf venation in Pilea peperomioides is a Voronoi diagram

CiCi Xingyu Zheng1, Shirsa Palit1, Matthew Venezia1, Elijah Blum1, Ullas V. Pedmale1, Dave Jackson1, Enrico Scarpella2, Przemyslaw Prusinkiewicz3, and Saket Navlakha1.
1Cold Spring Harbor Laboratory
2University of Alberta
3University of Calgary

Abstract

Reticulate leaf venation, characterized by the presence of loops, is a distinguishing feature of many flowering plants. However, our understanding of both the geometry and the morphogenesis of reticulate vein patterns is far from complete. We show that in the Chinese money plant (Pilea peperomioides), major veins form an approximate Voronoi diagram surrounding secretory pores known as hydathodes. We also propose a mechanistic model based on polar transport of the plant hormone auxin to produce Voronoi patterns, and we support this model with experimental evidence. In contrast to classical models where veins directly connect auxin sources to sinks, our model generates veins that bisect the space between adjacent auxin sources, collectively forming loops. The paradigm change offered by this model may open the door to study reticulate vein formation in other species.

Reference

CiCi Xingyu Zheng, Shirsa Palit, Matthew Venezia, Elijah Blum, Ullas V. Pedmale, Dave Jackson, Enrico Scarpella, Przemyslaw Prusinkiewicz, and Saket Navlakha. Reticulate leaf venation in Pilea peperomioides is a Voronoi diagram. Nature Communications (2026) 17:4111. https://doi.org/10.1038/s41467-026-71768-3.

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Movie 1
Simulation of auxin propagation in the absence of caps on auxin concentration and the rate of endocytosis with 2 auxin sources on a rectangluar grid (corresponds to Supplementary Figure 8).

Movie 2
Simulation of auxin propagation in the presence of a cap on auxin concentration with 2 auxin sources on a rectangluar grid (corresponds to Supplmentary Figure 9).

Movie 3
Simulation of auxin propagation in the presence of caps on auxin concentration and the rate of endocytosis with 2 auxin sources on a rectangular grid (corresponds to Fig. 4d and Supplementary Figure 10).

Movie 4
Simulation of auxin propagation in the presence of caps on auxin concentration and the rate of endocytosis with 5 auxin sources on a square grid (corresponds to Fig. 4e).

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Movie 9
Simulation of auxin driven patterning on Pilea leaves assuming auxin sources at hydathode positions. Movie 5 corresponds to the leaf sample shown in Fig. 5a. Movies 6-9 correspond to the leaf samples shown in Supplementary Figure 11a-d.

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Movie 13
Simulation of model progression in Supplementary Figure 14b, d-f.

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Simulation of improved model in Fig. 7b-e. Movies 16, 18, and 20 are zoomed-out versions of Movies 15, 17, and 19.