Sapala, A., Runions, A., Routier-Kierzkowska, A.-L., Das Gupta, M., Hong, L., Hofhuis, H., Verger, S., Mosca, G., Li, C.-B., Hay, A., Hamant, O., Roeder, A., Tsiantis, M., Prusinkiewicz, R., & Smith, R. S. (2018). Why plants make puzzle cells, and how their shape emerges. eLife, 7: e32794. doi:10.7554/eLife.32794.


Hervieux, N., Tsugawa, S., Fruleux, A., Dumond, M., Routier-Kierzkowska, A.-L., Komatsuzaki, T., Boudaoud, A., Larkin, J. C., Smith, R. S., Li, C.-B., & Hamant, O. (2017). Mechanical Shielding of Rapidly Growing Cells Buffers Growth Heterogeneity and Contributes to Organ Shape Reproducibility. Current Biology, 27(22), 3468-3479.e4. doi:10.1016/j.cub.2017.10.033.

Jones, A.R., Forero-Vargas, M., Withers, S.P., Smith, R.S., Traas, J., Dewitte, W., Murray, J.A.H. Cell-size dependent progression of the cell cycle creates homeostasis and flexibility of plant cell size. Nature Communications 8 (2017).

McKim, S. M., Routier-Kierzkowska, A.-L., Monniaux, M., Kierzkowski, D., Pieper, B., Smith, R. S., Tsiantis, M., & Hay, A. (2017). Seasonal Regulation of Petal Number. Plant Physiology, 175(2), 886-903. doi:10.1104/pp.17.00563.

Mosca, M., Sapala, A., Strauss, A., Routier-Kierzkowskai, A-L., Smith, R.S. On the micro-indentation of plant cells in a tissue context. Physical Biology 14 (2017).

Stamm, P., Strauß, S., Montenegro-Johnson, T. D., Smith, R. S., & Bassel, G. W. (2017). In Silico Methods for Cell Annotation, Quantification of Gene Expression, and Cell Geometry at Single-Cell Resolution Using 3DCellAtlas. PLANT HORMONES: METHODS AND PROTOCOLS, 3RD EDITION, 99-123. doi:10.1007/978-1-4939-6469-7_11.

Tsugawa, S., Hervieux, N., Kierzkowski, D., Routier-Kierzkowska, A.-L., Sapala, A., Hamant, O., Smith, R. S., Roeder, A. H. K., Boudaoud, A., & Li, C.-B. (2017). Clones of cells switch from reduction to enhancement of size variability in Arabidopsis sepals. Development, 144(23), 4398-4405. doi:10.1242/dev.153999.


Hong, L., Dumond, M., Tsugawa, S., Sapala, A., Routier-Kierzkowska, A-L., Zhou, Y., Chen, C., Kiss, A., Zhu, M., Hamant, O., Smith, R.S., Komatsuzaki, T., Li, C-B., Boudaoud, A., Roeder, A.H.K. Variable Cell Growth Yields Reproducible Organ Development through Spatiotemporal Averaging. Developmental Cell 38(1), 15-32 (2016).

Žádníková, P., Wabnik, K., Abuzeineh, A., Gallemi, M., Van Der Straeten, M., Smith, R.S., Inzé, D., Friml, J., Prusinkiewicz, P., Benková, E. A Model of Differential Growth-Guided Apical Hook Formation in Plants. The Plant Cell 28(10), 2464-2477 (2016)

Hofhuis, H., Moulton, D., Lessinnes, T., Routier-Kierzkowska, A.L., Bomphrey, R.J., Mosca, G., Reinhardt, H., Sarchet, P., Gan, X., Tsiantis, M. and Ventikos, Y. Morphomechanical Innovation Drives Explosive Seed Dispersal. Cell, 166: 222-233

Tsugawa, S., Hervieux, N., Hamant, O., Boudaoud, A., Smith, R. S., Li, C. B., and Komatsuzaki, T. Extracting Subcellular Fibrillar Alignment with Error Estimation: Application to Microtubules. Biophysical journal, 110: 1836-1844.

Hervieux, N., Dumond, M., Sapala, A., Routier-Kierzkowska, A.L., Kierzkowski, D., Roeder, A.H., Smith, R.S., Boudaoud, A. and Hamant, O. A mechanical feedback restricts sepal growth and shape in Arabidopsis. Current Biology, 26: 1019-1028.

Bassel, George W., and Richard S. Smith. Quantifying morphogenesis in plants in 4D. Current opinion in plant biology, 29: 87-94.

Kirchhelle, C., Chow, C.M., Foucart, C., Neto, H., Stierhof, Y.D., Kalde, M., Walton, C., Fricker, M., Smith, R.S., Jérusalem, A. and Irani, N. The Specification of Geometric Edges by a Plant Rab GTPase Is an Essential Cell-Patterning Principle During Organogenesis in Arabidopsis. Developmental cell, 36: 386-400.

von Wangenheim, D., Fangerau, J., Schmitz, A., Smith, R. S., Leitte, H., Stelzer, E. H., & Maizel, A. Rules and self-organizing properties of post-embryonic plant organ cell division patterns. Current Biology, 26: 439-449

Scheuring, D., Löfke, C., Krüger, F., Kittelmann, M., Eisa, A., Hughes, L., Smith, R.S., Hawes, C., Schumacher, K. and Kleine-Vehn, J. Actin-dependent vacuolar occupancy of the cell determines auxin-induced growth repression. Proceedings of the National Academy of Sciences, 113: 452-457.

Martinez, C. C., Chitwood, D. H., Smith, R. S., and Sinha, N. R. Left-right leaf asymmetry in decussate and distichous phyllotactic systems. BioRxiv, 043869.



Tauriello, G., Meyer, H. M., Smith, R. S., Koumoutsakos, P., and Roeder, A. H. Variability and constancy in cellular growth of Arabidopsis sepals. Plant physiology, 169: 2342-2358.

Weber, A., Braybrook, S., Huflejt, M., Mosca, G., Routier-Kierzkowska, A. L., and Smith, R. S. Measuring the mechanical properties of plant cells by combining micro-indentation with osmotic treatments. Journal of experimental botany, 66: 3229-3241.

Barbier de Reuille, P.B., Routier-Kierzkowska, A.L., Kierzkowski, D., Bassel, G.W., Schüpbach, T., Tauriello, G., Bajpai, N., Strauss, S., Weber, A., Kiss, A. and Burian, A., 2015. MorphoGraphX: a platform for quantifying morphogenesis in 4D. Elife, 4, e05864.

Montenegro-Johnson, T.D., Stamm, P., Strauss, S., Topham, A.T., Tsagris, M., Wood, A.T., Smith, R.S. and Bassel, G.W. Digital single-cell analysis of plant organ development using 3DCellAtlas.


Bassel GW, Stamm P, Mosca G, Barbier de Reuille P, Gibbs DJ, Winter R, Janka A, Holdsworth MJ, Smith RS*. Mechanical constraints imposed by 3D cellular geometry and arrangement modulate growth patterns in the Arabidopsis embryo. Proceedings of the National Academy of Science of USA. 111: 8685-8690.

Yoshida S, Barbier de Reuille P, Lane B, Bassel GW, Prusinkiewicz P, Smith RS*, Weijers D. Genetic control of plant development by overriding a geometric division rule. Developmental Cell, 29:75-87.

Vlad D, Kierzkowski D, Rast MI, Vuolo F, Dello Ioio R, Galinha C, Gan X, Hajheidari M, Hay A, Smith RS, Huijser P, Bailey CD, Tsiantis M. Leaf Shape Evolution Through Duplication, Regulatory Diversification, and Loss of a Homeobox Gene. Science, 343 (6172), 780-783.

Barbier de Reuille P, Robinson S, Smith RS. Quantifying Cell Shape and Gene Expression in the Shoot Apical Meristem Using MorphoGraphX. Plant Cell Morphogenesis, 121-134

Routier-Kierzkowska AL, Smith RS. Mechanical Measurements on Living Plant Cells by Micro-indentation with Cellular Force Microscopy. Plant Cell Morphogenesis, 135-146.

Smith RS. Modeling Plant Morphogenesis and Growth, in In “New Trends in the Physics and Mechanics of Biological Systems: Lecture Notes of the Les Houches Summer School: Volume 92, July 2009”, Oxford University Press 2011.


Kierzkowski D, Lenhard M, Smith RS, Kuhlemeier C. Interaction between Meristem Tissue Layers Controls Phyllotaxis. Developmental Cell, 26(6): 616-628.

Wabnik K, Robert HS, Smith RS, Friml J. Modeling Framework for the Establishment of the Apical-Basal Embryonic Axis in Plants. Current Biology 23:1-6.

Bert De Rybel B, Möller B, Yoshida S, Grabowicz I, Barbier de Reuille P, Boeren S, Smith RS, Borst JW, Weijers D. A bHLH Complex Controls Embryonic Vascular Tissue Establishment and Indeterminate Growth in Arabidopsis. Developmental Cell. 24(4): 426-437.

Routier-Kierzkowska AL, Smith RS*. Measuring the mechanics of morphogenesis. Current Opinion in Plant Biology. 16(1): 25-32.

Vogler H, Draeger C, Weber A, Felekis D, Eichenberger C, Routier‐Kierzkowska AL, Boisson‐Dernier A, Ringli C, Nelson BJ*, Smith RS*, Grossniklaus U*. The pollen tube: a soft shell with a hard core. The Plant Journal. 73(4): 617-627.


Routier-Kierzkowska AL; Smith RS*. Measuring the mechanics of morphogenesis. Current Opinion in Plant Biology. 16(1): 25-32.

Vogler H; Draeger C; Weber A; Felekis D; Eichenberger C; Routier‐Kierzkowska AL; Boisson‐Dernier A; Ringli C; Nelson BJ*; Smith RS*; Grossniklaus U*. The pollen tube: a soft shell with a hard core. The Plant Journal. 73(4): 617-627.

Guenot B; Bayer E; Kierzkowski D; Smith RS; Mandel T; Žádníková P; Benková E; Kuhlemeier C. Pin1-independent leaf initiation in Arabidopsis. Plant Phys. 159(4):1501-10.

Nakayama N; Smith RS; Mandel T; Kimura S; Boudaoud A; Kuhlemeier C.  Mechanical regulation of auxin-mediated growth. Current Biology 22(16):1468-76.

Aegerter-Wilmsen T; Heimlicher MB; Smith AC; Barbier de Reuille P; Smith RS; Aegerter CM; Basler K. Integrating Force Sensing and Signaling Pathways in a Model for the Regulation of Wing Imaginal Disc Size. Development 139(17):3221-3.

Felekis D; Weber A; Muntwyler S; Routier-Kierzkowska AL; Beyeler F; Smith RS; Nelson BJ. Automated stiffness characterization of living tobacco BY2 cells using the Cellular Force Microscope. Proceedings of the 8th IEEE International Conference on Automation Science and Engineering, 286-290.

Chitwood DH; Headland LR; Ranjan A; Braybrook SA; Koenig DP; Martinez C; Kuhlemeier C;  Smith RS; Sinha NR. Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning. Plant Cell 24(6):2318-27.

Kierzkowski D; Nakayama N; Routier-Kierzkowska AL; Weber A; Bayer E; Schorderet M; Reinhardt D; Kuhlemeier C; Smith RS*. Elastic domains regulate growth and organogenesis in the plant shoot apical meristem, Science 335:1096-1099.

 Routier-Kierzkowska AL; Weber A; Kochova P; Felekis D; Nelson B; Kuhlemeier C; Smith RS*.  Cellular Force Microscopy for in vivo Measurements of Plant tissue Mechanics. Plant Physiology, 158(4): 1514-1522.



Santuari L; Scacchi E; Rodriguez-Villalon A; Salinas P; Dohmann EMN; Brunoud G; Vernoux T; Smith RS; Hardtke CS.  Positional Information by Differential Endocytosis Splits Auxin Response to Drive Arabidopsis Root Meristem Growth. Current Biology 21:1-6.



Laňková M; Smith RS; Pešek B; Kubeš M; Zažímalová E; Petrášek J; Hoyerová K. Auxin influx inhibitors 1-NOA, 2-NOA, and CHPAA interfere with membrane dynamics in tobacco cells. Journal of Experimental Botany 61 (13): 3589-3598



Prusinkiewicz P; Crawford S; Smith RS; Ljung K; Bennett T; Ongaro V; Leyser O: Control of bud activation by an auxin transport switch, Proceedings of the National Academy of Science of USA. 106(41) 17431-17436.

Smith RS; Bayer EM. Auxin transport-feedback models of patterning in plants. Plant Cell and Environment, 32(9) 1258.

Bayer EM; Smith RS; Mandel T; Nakayama N; Sauer M; Prusinkiewicz P; and Kuhlemeier C. Integration of transport-based models for phyllotaxis and midvein formation. Genes and Development, 23:373-384.



Smith RS. The Role of Auxin Transport in Plant Patterning Mechanisms (primer). PLoS Biology, 6(12).



Smith RS; Kuhlemeier C; Prusinkiewicz P. Inhibition fields for phyllotactic pattern formation: a simulation study. Canadian Journal of Botany, 84(11).

Smith RS; Guyomarc’h S.; Mandel T; Reinhardt D; Kuhlemeier C; Prusinkiewicz P. A plausible model of phyllotaxis. Proceedings of the National Academy of Science of USA. 103: 1301–1306.

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