Mosca G, Eng RC, Adibi M, Yoshida S, Lane B, Bergheim L, Weber G, Smith RS, Hay A: Growth and tension in explosive fruit. Current Biology 2024,

Emonet A, Perez-Anton M, Neumann U, Dunemann S, Huettel B, Koller R, Hay A: Amphicarpic development in the emerging model organism Cardamine chenopodiifolia. bioRxiv 2024,

Emonet A, Awad M, Tikhomirov N, Vasilarou M, Perez-Anton M, Gan X, Novikova P, Hay A: Polyploid genome assembly of Cardamine chenopodiifolia. bioRxiv 2024,



Pérez-Antón, M., Schneider, I., Kroll, P., Hofhuis, H., Metzger, S., Pauly, M., Hay, A. (2022) Explosive seed dispersal depends on SPL7 to ensure sufficient copper for localized lignin deposition via laccases. PNAS 119: e2202287119,

Emonet. A, Hay, A. (2022) Development and diversity of lignin patterns. Plant Physiology 190: 31–43,



Galstyan, A., Sarchet, P., Campos-Martin, R., Adibi, M., Nikolov, L.A., Pérez Antón, M., Hofhuis, H., Rambaud-Lavigne, L., Gan, X., Hay, A. (2021) INDEHISCENT regulates explosive seed dispersal. bioRxiv,

Akiyama, R., Sun, J., Hatakeyama, M., Lischer, H.E.L., Briskine, R.V., Hay, A., Gan, X., Tsiantis, M., Kudoh, H., Kanaoka, M.M, Sese, J., Shimizu, K.K., Shimizu?Inatsugi, R. (2021) Fine?scale empirical data on niche divergence and homeolog expression patterns in an allopolyploid and its diploid progenitor species. New Phytologist 229: 3587–3601,



Rambaud-Lavigne, L., Hay, A. (2020) Floral organ development goes live. Journal of Experimental Botany 71: 2472–8, doi:10.1093/jxb/eraa038

Pérez-Antón, M., Hay, A. (2020) Schooling PhD students in plant development. New Phytologist 226:1544–7,

Sun, J., Shimizu-Inatsugi, R., Hofhuis, H., Shimizu, K., Hay, A., Shimizu, K.K., Sese, J. (2021) A recently formed triploid Cardamine insueta inherits leaf vivipary and submergence tolerance traits of parents. Frontiers in Genetics 11: 1067,



Neumann, U., Hay, A. (2019) Seed coat development in explosively dispersed seeds of Cardamine hirsuta. Annals of Botany 126: 39–59, doi: 10.1093/aob/mcz190



Galstyan, A; Hay, A (2018) Snap, crack and pop of explosive fruit. Current opinion in genetics & development 51, 31-36

Vuolo, F., Kierzkowski, D., Runions, A., Hajheidari, M., Mentink, R. A., Das Gupta, M., Zhang, Z., Vlad, D., Wang, Y., Pecinka, A., Gan, X., Hay, A., Huijser, P., & Tsiantis, M. (2018). LMI1 homeodomain protein regulates organ proportions by spatial modulation of endoreduplication. Genes and Development. doi:10.1101/gad.318212.118.

Monniaux, M., Pieper, B., McKim, S. M., Routier-Kierzkowska, A.-L., Kierzkowski, D., Smith, R. S., & Hay, A. (2018). The role of APETALA1 in petal number robustness. eLife, 7: e39399. doi:10.7554/eLife.39399.

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.



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, 886-903. doi:10.1104/pp.17.00563.

Hofhuis, H. and Hay, A. (2017), Explosive seed dispersal. New Phytologist 216, 339-342. doi:10.1111/nph.14541.

Monniaux, M., McKim, S. M., Cartolano, M., Thévenon, E., Parcy, F., Tsiantis, M. and Hay, A. (2017), Conservation vs divergence in LEAFY and APETALA1 functions between Arabidopsis thaliana and Cardamine hirsuta. New Phytologist 216, 549-561, doi:10.1111/nph.14419.



Monniaux M, Hay A. Cells, Walls, and Endless Forms. Curr Opin Plant Biol 34:114-121 (2016).

Gan, X.*, Hay, A.*, Kwantes, M.*, Haberer, G., Hallab, A., Dello Ioio, R., Hofhuis, H., Pieper, B., Cartolano, M., Neumann, U., Nikolov, L. A., Song, B., Hajheidari, M., Briskine, R., Kougioumoutzi, E., Vlad, D., Broholm, S., Hein, J., Meksem, K., Lightfoot, D., Shimizu, K. K., Shimizu-Inatsugi, R., Imprialou, M., Kudrna, D., Wing, R., Sato, S., Huijser, P., Filatov, D., Mayer, K. F. X., Mott, R., Tsiantis, M. The Cardamine hirsuta genome offers insight into the evolution of morphological diversity. Nature Plants, 2, 16167, DOI: 10.1038/nplants.2016.167 (2016). * contributed equally.

Hay A, Tsiantis M. Cardamine hirsuta: a comparative view. Curr Opin Genet Dev 39:1-7 (2016).

Hofhuis, H. Moulton, D., Lessinnes, T., Routier-Kierzkowska, A.-L., Bomphrey, R.J., Mosca, G., Reinhardt, H., Sarchet, P., Gan, X., Tsiantis, M., Ventikos, Y., Walker, S, Goriely, A., Smith, R. and A. Hay: Morphomechanical Innovation Drives Explosive Seed Dispersal. Cell 166, 1, 222-2333, DOI: 10.1016/j.cell.2016.05.002 (2016).

Pieper B, Monniaux M, Hay A. The genetic architecture of petal number in Cardamine hirsuta. New Phytologist 209(1):395-406. doi: 10.1111/nph.13586 (2016)



Rast-Somssich, M.; Broholm, S.; Jenkins, H.; Canales, C.; Vlad, D.; Kwantes, M.; Bilsborough, G.; Dello Ioio, R.; Ewing, R. M.; Laufs, P.; Huijser, P.; Ohno, C.; Heisler, M. G.; Hay, A.; Tsiantis, M. (2015) Alternate wiring of a KNOXI genetic network underlies differences in leaf development of A. thaliana and C. hirsute. Genes and Development 29: 2391-2404.

Monniaux M, Pieper B, Hay A. (2015). Stochastic variation in Cardamine hirsuta petal number. Annals of Botany doi:10.1093/aob/mcv131

Cartolano M, Pieper B, Lempe J, Tattersall A, Huijser P, Tresch A, Darrah PR, Hay A, Tsiantis M. (2015). Heterochrony underpins natural variation in Cardamine hirsuta leaf form. PNAS 112(33): 10539-44

Barbier de Reuille P, Routier-Kierzkowska A, Kierzkowski D, Bassel GW, Schüpbach T, Tauriello G, Bajpai N, Strauss S, Weber A, Kiss A, Burian A, Hofhuis H, Sapala A, Lipowczan M, Heimlicher MB, Robinson S, Bayer EM, Basler K, Koumoutsakos P, Roeder A, Aegerter-Wilmsen T, Nakayama N, Tsiantis M, Hay A, Kwiatkowska D, Xenarios I, Kuhlemeier C, Smith RS (2015) MorphoGraphX: A platform for quantifying morphogenesis in 4D. eLife 4:e05864



Lewis MW, Bolduc N, Hake K, Htike Y, Hay A, Candela H, Hake S (2014) Gene regulatory interactions at lateral organ boundaries in maize. Development 141(23): 4590-7

Vlad D, Kierzkowski D, Rast M I, Vuolo F, Dello Ioio R, Galinha C, Gan X, Hajheidari M, Hay A, Smith R S, Huijser P, Bailey C D, Tsiantis M (2014). Leaf shape evolution through duplication, regulatory diversification and loss of a homeobox gene. Science 343: 780-3

Hay A*, Pieper B, Cooke E, Mandáková T, Cartolano M, Tattersall A, Dello Ioio R, McGowan S, Barkoulas M, Galinha C, Rast M, Hofhuis H, Then C, Plieske J, Ganal M, Mott R, Martinez-Garcia J, Carine M, Scotland R, Gan X, Filatov D, Lysak M, Tsiantis M* (2014). Cardamine hirsuta: a versatile genetic system for comparative studies. The Plant Journal 78(1): 1-15, *joint corresponding authors



Kougioumoutzi E, Cartolano M, Canales C, Dupré M, Bramsiepe J, Vlad D, Rast M, Dello Ioio R, Tattersall A, Schnittger A, Hay A*, Tsiantis M.* (2013). SIMPLE LEAF3 encodes a ribosome-associated protein required for leaflet development in Cardamine hirsuta. The Plant Journal 73(4): 533-45, *joint corresponding authors



Bilsborough, G., Runions, A., Barkoulas, M., Jenkins, H., Hasson, A., Galinha, C., Laufs, P., Hay, A., Prusinkiewicz, P. and Tsiantis, M. (2011) A model for the regulation of Arabidopsis thaliana leaf margin development. PNAS 108(8), 3424-9



McKim, S and Hay, A. (2010) Patterning and evolution of floral structures – marking time. Current Opinion in Genetics & Development 20(4): 448-53

Piazza, P., Bailey, C.D., Cartolano, M., Krieger, J., Cao, J., Ossowski, S., Schneeberger, K., Hall, N., Fei He, Juliette de Meaux, MacLeod, N., Filatov D., Hay, A.* and Tsiantis, M.* (2010) Arabidopsis thaliana leaf form evolved via loss of KNOX expression in leaves in association with a selective sweep. Current Biology 20(24): 2223-8 *co-corresponding authors

Hay, A. and Tsiantis, M. (2010) KNOX genes: versatile regulators of plant development and diversity. Development 137, 3153-65



Hay, A. and Tsiantis, M. (2009) A KNOX family TALE. Current Opinion in Plant Biology. 12(5): 593-8



Blein, T., Pulido, A., Vialette-Guiraud, A., Nikovics, K., Morin, H., Hay, A., Johansen, I.E., Tsiantis, M. and Laufs, P. (2008). A conserved molecular framework for compound leaf development. Science 322: 1835-8

Nature News (18 December 2008) doi:10.1038/news.2008.1316

Faculty of 1000 Biology:

Jasinski, S., Tattersall, A., Piazza, P., Hay, A., Martinez-Garcia, J., Schmitz, G., Theres, K., McCormick, S., Tsiantis, M. (2008). PROCERA encodes a DELLA protein that mediates control of dissected leaf form in tomato. The Plant Journal 56: 603-12

Barkoulas, M., Hay, A., Kougioumoutzi E, and Tsiantis, M. (2008). A developmental framework for dissected leaf formation in the Arabidopsis relative Cardamine hirsuta. Nature Genetics 40: 1136-41

Faculty of 1000 Biology:



Hay A, Barkoulas M, Tsiantis M (2006). ASYMMETRIC LEAVES1 and auxin activities converge to repress BREVIPEDICELLUS expression and promote leaf development in Arabidopsis. Development 133(20): 3955-61.

In this Issue (2006). Development 133(20): 2001.

Hay, A. and Tsiantis, M. (2006). The genetic basis for differences in leaf form between Arabidopsis thaliana and its wild relative Cardamine hirsuta. Nature Genetics 38: 942-7.

Editor’s Choice (2006). Science 313: 411

News & Views (2006). Nature Genetics 38: 865-6

Faculty of 1000 Biology:



Hay, A. and Tsiantis, M. (2005). From genes to plants via meristems. Development 132: 2679-84

Grigg, S., Canales, C., Hay, A. and Tsiantis, M. (2005). SERRATE coordinates shoot meristem function and leaf axial patterning in Arabidopsis. Nature 437: 1022-6

Jasinski, S., Piazza, P. Craft, J., Hay, A., Woolley, L., Rieu, I., Phillips, A., Hedden, P. and Tsiantis, M. (2005). KNOX action in Arabidopsis is mediated by coordinate regulation of cytokinin and gibberellin activities. Current Biology 15: 1560-5

Faculty of 1000 Biology:



Foster, T.*, Hay, A.*, Johnston, R. and Hake, S. (2004). The establishment of axial patterning in the maize leaf. Development 131(16): 3921-9 *co-first authors.

Hay, A. and Hake, S. (2004). The dominant mutant Wavy auricle in blade1 ( Wab1) disrupts patterning in a lateral domain of the maize leaf. Plant Physiology 135(1): 300-8

Hay A, Barkoulas M, Tsiantis M. (2004). PINning down the connections: transcription factors and hormones in leaf morphogenesis. Current Opinion in Plant Biology 7: 575-81

Hay, A., Craft, J. and Tsiantis, M. (2004). Hormones and Homeoboxes. Bridging the Gap? Bioessays 26(4): 395-404.



Hay, A., Jackson, D., Ori, N. and Hake, S. (2003). Analysis of the Competence to Respond to KNOTTED1 Activity in Arabidopsis Leaves Using a Steroid Induction System. Plant Physiology 131: 1671-80

Tsiantis, M. and Hay, A. (2003). Comparative plant development: the time of the leaf? Nature Reviews Genetics 4: 169-80



Hay, A., Kaur, H., Phillips, A., Hedden, P., Hake, S. and Tsiantis, M. (2002). The gibberellin pathway mediates KNOTTED1-type homeobox function in plants with different body plans. Current Biology 12(20): 1557-65

Dispatch (2006). Current Biology 12(20): R696-8

Faculty of 1000 Biology:

Kaplinsky N, Braun D, Lisch D, Hay A, Hake S, Freeling M (2002) Biodiversity (Communications arising): maize transgene results in Mexico are artefacts. Nature 416: 601-02; discussion 600, 602

Tsiantis M, Hay A, Ori N, Kaur H, Henderson I, Holtan H, McCormick S, Hake S (2002) Developmental signals regulating leaf form. In Developmental Genetics and Plant Evolution. Edited by Cronk QCB, Bateman RM, Hawkins JA: Taylor & Francis; 418-430. [Warren A (Series Editor): The Systematics Association Special Volume Series, vol 65.]



Hay, A., Watson, L., Zhang, C. and McManus, M. (1998). Identification of cell wall proteins in roots of phosphate-deprived white clover plants. Plant Physiol. Biochem. 36(4): 305-11

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