{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,27]],"date-time":"2026-10-27T21:52:59Z","timestamp":1793137979813,"version":"4.5.0"},"reference-count":94,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T00:00:00Z","timestamp":1787616000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T00:00:00Z","timestamp":1787616000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["Cluster of Excellence CEPLAS (EXC 2048\/1 Project ID: 390686111)"],"award-info":[{"award-number":["Cluster of Excellence CEPLAS (EXC 2048\/1 Project ID: 390686111)"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["CO 318\/14-1"],"award-info":[{"award-number":["CO 318\/14-1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["EXC 2048\/1 Project ID: 390686111"],"award-info":[{"award-number":["EXC 2048\/1 Project ID: 390686111"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004189","name":"Max-Planck-Gesellschaft","doi-asserted-by":"publisher","award":["Core Grant"],"award-info":[{"award-number":["Core Grant"]}],"id":[{"id":"10.13039\/501100004189","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004189","name":"Max-Planck-Gesellschaft","doi-asserted-by":"publisher","award":["Core Grant"],"award-info":[{"award-number":["Core Grant"]}],"id":[{"id":"10.13039\/501100004189","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004543","name":"China Scholarship Council","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100004543","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100005156","name":"Alexander von Humboldt-Stiftung","doi-asserted-by":"publisher","award":["ESP - 1238682 - HFST-P"],"award-info":[{"award-number":["ESP - 1238682 - HFST-P"]}],"id":[{"id":"10.13039\/100005156","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100005156","name":"Alexander von Humboldt-Stiftung","doi-asserted-by":"publisher","award":["ITA 1216206 HFST-P"],"award-info":[{"award-number":["ITA 1216206 HFST-P"]}],"id":[{"id":"10.13039\/100005156","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100010661","name":"EC | Horizon 2020 Framework Programme","doi-asserted-by":"publisher","award":["HORIZON-MSCA-2023-PF-01"],"award-info":[{"award-number":["HORIZON-MSCA-2023-PF-01"]}],"id":[{"id":"10.13039\/100010661","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Commun"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Developmental transitions occur in the life cycles of all multicellular organisms. Despite their fundamental relevance, the underlying dynamics remain poorly understood. In plants, floral transition is a key developmental process whereby the shoot apical meristem changes from producing leaves to forming flowers. Using quantitative imaging, developmental genetics, and dynamical systems theory, we show that a time-dependent bistable switch between expression of\n                    <jats:italic>APETALA2<\/jats:italic>\n                    , a key floral inhibitor, and the floral activators\n                    <jats:italic>SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1<\/jats:italic>\n                    and\n                    <jats:italic>FRUITFULL<\/jats:italic>\n                    , can explain the dynamics of floral transition in Arabidopsis. Notably, we detect a slowing down of the inhibitor dynamics, consistent with the system crossing a critical point of a bistable switch and transiently experiencing a ghost attractor. We demonstrate that this time-dependent bistability is essential to generate the range of dynamical behaviours measured across genotypes, including oscillations in the inhibitor, and that it also confers robustness in the transition. Collectively, our work provides quantitative evidence of time-dependent bistability underlying floral transition, which introduces a new timescale to this developmental process.\n                  <\/jats:p>","DOI":"10.1038\/s41467-026-76210-2","type":"journal-article","created":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T12:44:20Z","timestamp":1787661860000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Time-dependent bistability leads to critical slowing down during floral transition in Arabidopsis"],"prefix":"10.1038","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8926-2709","authenticated-orcid":false,"given":"Gabriel","family":"Rodr\u00edguez-Maroto","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-6177-2730","authenticated-orcid":false,"given":"Kang","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5445-7469","authenticated-orcid":false,"given":"Pau","family":"Casanova-Ferrer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9654-252X","authenticated-orcid":false,"given":"Martina","family":"Cerise","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6988-4172","authenticated-orcid":false,"given":"George","family":"Coupland","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3005-597X","authenticated-orcid":false,"given":"Pau","family":"Formosa-Jordan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,8,25]]},"reference":[{"key":"76210_CR1","doi-asserted-by":"publisher","first-page":"20220002","DOI":"10.1098\/rsfs.2022.0002","volume":"12","author":"M S\u00e1ez","year":"2022","unstructured":"S\u00e1ez, M., Briscoe, J. & Rand, D. A. Dynamical landscapes of cell fate decisions. Interface Focus 12, 20220002 (2022).","journal-title":"Interface Focus"},{"key":"76210_CR2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-28151-9","volume":"13","author":"P-F Lenne","year":"2022","unstructured":"Lenne, P.-F. & Trivedi, V. Sculpting tissues by phase transitions. Nat. Commun. 13, 664 (2022).","journal-title":"Nat. Commun."},{"key":"76210_CR3","doi-asserted-by":"publisher","DOI":"10.1016\/j.heliyon.2025.e42995","volume":"11","author":"E Force","year":"2025","unstructured":"Force, E., Lamy, D., Debernard, S., Savour\u00e9, A. & Dacher, M. Developmental transitions involve common biological processes across living beings. Heliyon 11, e42995 (2025).","journal-title":"Heliyon"},{"key":"76210_CR4","unstructured":"Waddington, C. H. The Strategy of the Genes. A Discussion of Some Aspects of Theoretical Biology. (Routledge, London and New York, 1957)."},{"key":"76210_CR5","doi-asserted-by":"publisher","DOI":"10.1186\/1752-0509-8-43","volume":"8","author":"B Verd","year":"2014","unstructured":"Verd, B., Crombach, A. & Jaeger, J. Classification of transient behaviours in a time-dependent toggle switch model. BMC Syst. Biol. 8, 43 (2014).","journal-title":"BMC Syst. Biol."},{"key":"76210_CR6","doi-asserted-by":"crossref","unstructured":"Jaeger, J. & Crombach, A. Life\u2019s attractors. in Evolutionary Systems Biology, Adv. Exp. Med. Biol. 751, 93\u2013119 (Springer, New York, 2012).","DOI":"10.1007\/978-1-4614-3567-9_5"},{"key":"76210_CR7","doi-asserted-by":"publisher","first-page":"R458","DOI":"10.1016\/j.cub.2012.03.045","volume":"22","author":"JE Ferrell","year":"2012","unstructured":"Ferrell, J. E. Bistability, bifurcations, and Waddington\u2019s epigenetic landscape. Curr. Biol. 22, R458\u2013R466 (2012).","journal-title":"Curr. Biol."},{"key":"76210_CR8","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1002\/bies.201100031","volume":"34","author":"S Huang","year":"2012","unstructured":"Huang, S. The molecular and mathematical basis of Waddington\u2019s epigenetic landscape: a framework for post-Darwinian biology? BioEssays 34, 149\u2013157 (2012).","journal-title":"BioEssays"},{"key":"76210_CR9","doi-asserted-by":"publisher","first-page":"140","DOI":"10.1016\/S0955-0674(02)00314-9","volume":"14","author":"JE Ferrell","year":"2002","unstructured":"Ferrell, J. E. Self-perpetuating states in signal transduction: positive feedback, double-negative feedback and bistability. Curr. Opin. Cell Biol. 14, 140\u2013148 (2002).","journal-title":"Curr. Opin. Cell Biol."},{"key":"76210_CR10","doi-asserted-by":"publisher","first-page":"e1008231","DOI":"10.1371\/journal.pcbi.1008231","volume":"17","author":"J Rombouts","year":"2021","unstructured":"Rombouts, J. & Gelens, L. Dynamic bistable switches enhance robustness and accuracy of cell cycle transitions. PLoS Comput. Biol. 17, e1008231 (2021).","journal-title":"PLoS Comput. Biol."},{"key":"76210_CR11","doi-asserted-by":"crossref","unstructured":"Strogatz, S. H. Nonlinear Dynamics and Chaos: With Applications to Physics, Biology, Chemistry, and Engineering (Chapman and Hall\/CRC, 2018).","DOI":"10.1201\/9780429399640"},{"key":"76210_CR12","doi-asserted-by":"publisher","DOI":"10.1242\/dev.204617","volume":"152","author":"U Kadiyala","year":"2025","unstructured":"Kadiyala, U. et al. From genes to patterns: five key dynamical systems concepts to decode developmental regulatory mechanisms. Development 152, dev204617 (2025).","journal-title":"Development"},{"key":"76210_CR13","doi-asserted-by":"publisher","first-page":"10501","DOI":"10.1103\/PhysRevB.40.10501","volume":"40","author":"SH Strogatz","year":"1989","unstructured":"Strogatz, S. H. & Westervelt, R. M. Predicted power laws for delayed switching of charge-density waves. Phys. Rev. B. 40, 10501\u201310508 (1989).","journal-title":"Phys. Rev. B."},{"key":"76210_CR14","doi-asserted-by":"publisher","first-page":"093064","DOI":"10.1088\/1367-2630\/abb549","volume":"22","author":"J Sardany\u00e9s","year":"2020","unstructured":"Sardany\u00e9s, J., Raich, C. & Alarc\u00f3n, T. Noise-induced stabilization of saddle-node ghosts. New J. Phys. 22, 093064 (2020).","journal-title":"New J. Phys."},{"key":"76210_CR15","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1016\/j.cels.2022.12.003","volume":"14","author":"O Karin","year":"2023","unstructured":"Karin, O., Miska, E. A. & Simons, B. D. Epigenetic inheritance of gene silencing is maintained by a self-tuning mechanism based on resource competition. Cell Syst. 14, 24\u201340.e11 (2023).","journal-title":"Cell Syst."},{"key":"76210_CR16","doi-asserted-by":"publisher","first-page":"047202","DOI":"10.1103\/PhysRevLett.133.047202","volume":"133","author":"D Koch","year":"2024","unstructured":"Koch, D. et al. Ghost channels and ghost cycles guiding long transients in dynamical systems. Phys. Rev. Lett. 133, 047202 (2024).","journal-title":"Phys. Rev. Lett."},{"key":"76210_CR17","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1016\/j.physd.2019.02.005","volume":"395","author":"JH Li","year":"2019","unstructured":"Li, J. H., Ye, F. X.-F., Qian, H. & Huang, S. Time-dependent saddle\u2013node bifurcation: breaking time and the point of no return in a non-autonomous model of critical transitions. Phys. Nonlinear Phenom. 395, 7\u201314 (2019).","journal-title":"Phys. Nonlinear Phenom."},{"key":"76210_CR18","doi-asserted-by":"publisher","first-page":"e1004097","DOI":"10.1371\/journal.pcbi.1004097","volume":"11","author":"C Meisel","year":"2015","unstructured":"Meisel, C., Klaus, A., Kuehn, C. & Plenz, D. Critical slowing down governs the transition to neuron spiking. PLoS Comput. Biol. 11, e1004097 (2015).","journal-title":"PLoS Comput. Biol."},{"key":"76210_CR19","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-15908-3","volume":"11","author":"MI Maturana","year":"2020","unstructured":"Maturana, M. I. et al. Critical slowing down as a biomarker for seizure susceptibility. Nat. Commun. 11, 2172 (2020).","journal-title":"Nat. Commun."},{"key":"76210_CR20","doi-asserted-by":"publisher","first-page":"3683","DOI":"10.1073\/pnas.1324186111","volume":"111","author":"D Krotov","year":"2014","unstructured":"Krotov, D., Dubuis, J. O., Gregor, T. & Bialek, W. Morphogenesis at criticality. Proc. Natl. Acad. Sci. USA. 111, 3683\u20133688 (2014).","journal-title":"Proc. Natl. Acad. Sci. USA."},{"key":"76210_CR21","doi-asserted-by":"publisher","first-page":"1724","DOI":"10.1016\/j.bpj.2015.08.024","volume":"109","author":"DE Tufcea","year":"2015","unstructured":"Tufcea, D. E. & Fran\u00e7ois, P. Critical timing without a timer for embryonic development. Biophys. J. 109, 1724\u20131734 (2015).","journal-title":"Biophys. J."},{"key":"76210_CR22","doi-asserted-by":"publisher","first-page":"17546","DOI":"10.1073\/pnas.1406326111","volume":"111","author":"V Dakos","year":"2014","unstructured":"Dakos, V. & Bascompte, J. Critical slowing down as early warning for the onset of collapse in mutualistic communities. Proc. Natl. Acad. Sci. USA 111, 17546\u201317551 (2014).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"76210_CR23","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1063\/1.1349894","volume":"11","author":"JE Ferrell","year":"2001","unstructured":"Ferrell, J. E. & Xiong, W. Bistability in cell signaling: how to make continuous processes discontinuous, and reversible processes irreversible. Chaos 11, 227\u2013236 (2001).","journal-title":"Chaos"},{"key":"76210_CR24","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1126\/science.1156951","volume":"321","author":"TYC Tsai","year":"2008","unstructured":"Tsai, T. Y. C. et al. Robust, tunable biological oscillations from interlinked positive and negative feedback loops. Science 321, 126\u2013129 (2008).","journal-title":"Science"},{"key":"76210_CR25","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1038\/35002131","volume":"403","author":"TS Gardner","year":"2000","unstructured":"Gardner, T. S., Cantor, C. R. & Collins, J. J. Construction of a genetic toggle switch in Escherichia coli. Nature 403, 339\u2013342 (2000).","journal-title":"Nature"},{"key":"76210_CR26","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.cels.2021.08.013","volume":"13","author":"M S\u00e1ez","year":"2022","unstructured":"S\u00e1ez, M. et al. Statistically derived geometrical landscapes capture principles of decision-making dynamics during cell fate transitions. Cell Syst. 13, 12\u201328.e3 (2022).","journal-title":"Cell Syst."},{"key":"76210_CR27","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1016\/j.ydbio.2007.02.036","volume":"305","author":"S Huang","year":"2007","unstructured":"Huang, S., Guo, Y. P., May, G. & Enver, T. Bifurcation dynamics in lineage-commitment in bipotent progenitor cells. Dev. Biol. 305, 695\u2013713 (2007).","journal-title":"Dev. Biol."},{"key":"76210_CR28","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1038\/s44320-025-00164-8","volume":"22","author":"BD Simons","year":"2025","unstructured":"Simons, B. D. & Karin, O. Cell cycle criticality as a mechanism for robust cell population control. Mol. Syst. Biol. 22, 241\u2013258 (2025).","journal-title":"Mol. Syst. Biol."},{"key":"76210_CR29","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1007\/BF02426835","volume":"6","author":"SF Gilbert","year":"1991","unstructured":"Gilbert, S. F. Epigenetic Landscaping: Waddington\u2019s Use of Cell Fate Bifurcation Diagrams. Biol. and Philos. 6, 135\u2013154 (1991).","journal-title":"Biol. and Philos."},{"key":"76210_CR30","doi-asserted-by":"publisher","first-page":"e3626","DOI":"10.1371\/journal.pone.0003626","volume":"3","author":"ER \u00c1lvarez-Buylla","year":"2008","unstructured":"\u00c1lvarez-Buylla, E. R. et al. Floral morphogenesis: stochastic explorations of a gene network epigenetic landscape. PLoS ONE 3, e3626 (2008).","journal-title":"PLoS ONE"},{"key":"76210_CR31","doi-asserted-by":"publisher","first-page":"2490","DOI":"10.1093\/jxb\/eraa057","volume":"71","author":"A Kinoshita","year":"2020","unstructured":"Kinoshita, A. & Richter, R. Genetic and molecular basis of floral induction in Arabidopsis thaliana. J. Exp. Bot. 71, 2490\u20132504 (2020).","journal-title":"J. Exp. Bot."},{"key":"76210_CR32","doi-asserted-by":"publisher","first-page":"160","DOI":"10.3389\/fgene.2015.00160","volume":"6","author":"J Davila-Velderrain","year":"2015","unstructured":"Davila-Velderrain, J., Martinez-Garcia, J. C. & Alvarez-Buylla, E. R. Modeling the epigenetic attractors landscape: toward a post-genomic mechanistic understanding of development. Front. Genet. 6, 160 (2015).","journal-title":"Front. Genet."},{"key":"76210_CR33","doi-asserted-by":"publisher","first-page":"2565","DOI":"10.1093\/jxb\/erw123","volume":"67","author":"J Davila-Velderrain","year":"2016","unstructured":"Davila-Velderrain, J., Martinez-Garcia, J. C. & Alvarez-Buylla, E. R. Dynamic network modelling to understand flowering transition and floral patterning. J. Exp. Bot. 67, 2565\u20132572 (2016).","journal-title":"J. Exp. Bot."},{"key":"76210_CR34","doi-asserted-by":"publisher","first-page":"192","DOI":"10.1126\/science.abg5999","volume":"373","author":"E Azpeitia","year":"2021","unstructured":"Azpeitia, E. et al. Cauliflower fractal forms arise from perturbations of floral gene networks. Science 373, 192\u2013197 (2021).","journal-title":"Science"},{"key":"76210_CR35","doi-asserted-by":"publisher","first-page":"852047","DOI":"10.3389\/fpls.2022.852047","volume":"13","author":"EC Ch\u00e1vez-Hern\u00e1ndez","year":"2022","unstructured":"Ch\u00e1vez-Hern\u00e1ndez, E. C., Quiroz, S., Garc\u00eda-Ponce, B. & \u00c1lvarez-Buylla, E. R. The flowering transition pathways converge into a complex gene regulatory network that underlies the phase changes of the shoot apical meristem in Arabidopsis thaliana. Front. Plant Sci. 13, 852047 (2022).","journal-title":"Front. Plant Sci."},{"key":"76210_CR36","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-46941-1","volume":"15","author":"E Coen","year":"2024","unstructured":"Coen, E. & Prusinkiewicz, P. Developmental timing in plants. Nat. Commun. 15, 2674 (2024).","journal-title":"Nat. Commun."},{"key":"76210_CR37","doi-asserted-by":"publisher","first-page":"655","DOI":"10.1016\/j.cell.2006.05.005","volume":"125","author":"I B\u00e4urle","year":"2006","unstructured":"B\u00e4urle, I. & Dean, C. The timing of developmental transitions in plants. Cell 125, 655\u2013664 (2006).","journal-title":"Cell"},{"key":"76210_CR38","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-18782-1","volume":"11","author":"Y Zhu","year":"2020","unstructured":"Zhu, Y. et al. TERMINAL FLOWER 1-FD complex target genes and competition with FLOWERING LOCUS T. Nat. Commun. 11, 5118 (2020).","journal-title":"Nat. Commun."},{"key":"76210_CR39","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1007\/s10265-020-01174-3","volume":"133","author":"M Fuchs","year":"2020","unstructured":"Fuchs, M. & Lohmann, J. U. Aiming for the top: non-cell autonomous control of shoot stem cells in Arabidopsis. J. Plant Res. 133, 297\u2013309 (2020).","journal-title":"J. Plant Res."},{"key":"76210_CR40","doi-asserted-by":"publisher","first-page":"e3001043","DOI":"10.1371\/journal.pbio.3001043","volume":"19","author":"DS \u00d3\u2019Maoil\u00e9idigh","year":"2021","unstructured":"\u00d3\u2019Maoil\u00e9idigh, D. S. et al. Systematic analyses of the MIR172 family members of Arabidopsis define their distinct roles in regulation of APETALA2 during floral transition. PLoS Biol. 19, e3001043 (2021).","journal-title":"PLoS Biol."},{"key":"76210_CR41","doi-asserted-by":"publisher","DOI":"10.1242\/dev.202089","volume":"150","author":"M Cerise","year":"2023","unstructured":"Cerise, M. et al. Two modes of gene regulation by TFL1 mediate its dual function in flowering time and shoot determinacy of Arabidopsis. Development 150, dev202089 (2023).","journal-title":"Development"},{"key":"76210_CR42","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.60661","volume":"9","author":"A Kinoshita","year":"2020","unstructured":"Kinoshita, A. et al. Regulation of shoot meristem shape by photoperiodic signaling and phytohormones during floral induction of Arabidopsis. eLife 9, e60661 (2020).","journal-title":"eLife"},{"key":"76210_CR43","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-51341-6","volume":"15","author":"E Bertran Garcia de Olalla","year":"2024","unstructured":"Bertran Garcia de Olalla, E. et al. Coordination of shoot apical meristem shape and identity by APETALA2 during floral transition in Arabidopsis. Nat. Commun. 15, 6930 (2024).","journal-title":"Nat. Commun."},{"key":"76210_CR44","doi-asserted-by":"publisher","first-page":"2156","DOI":"10.1105\/tpc.110.075606","volume":"22","author":"L Yant","year":"2010","unstructured":"Yant, L. et al. Orchestration of the floral transition and floral development in Arabidopsis by the bifunctional transcription factor APETALA2. Plant Cell 22, 2156\u20132170 (2010).","journal-title":"Plant Cell"},{"key":"76210_CR45","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1104\/pp.112.202614","volume":"160","author":"RGH Immink","year":"2012","unstructured":"Immink, R. G. H. et al. Characterization of SOC1\u2019s central role in flowering by the identification of its upstream and downstream regulators. Plant Physiol. 160, 433\u2013449 (2012).","journal-title":"Plant Physiol."},{"key":"76210_CR46","doi-asserted-by":"publisher","first-page":"107717","DOI":"10.1016\/j.celrep.2020.107717","volume":"31","author":"M Romera-Branchat","year":"2020","unstructured":"Romera-Branchat, M. et al. Functional divergence of the Arabidopsis Florigen-Interacting bZIP Transcription Factors FD and FDP. Cell Rep. 31, 107717 (2020).","journal-title":"Cell Rep."},{"key":"76210_CR47","doi-asserted-by":"publisher","first-page":"686","DOI":"10.1038\/s41586-025-09704-6","volume":"648","author":"H Gao","year":"2025","unstructured":"Gao, H. et al. Florigen activation complex forms via multifaceted assembly in Arabidopsis. Nature 648, 686\u2013695 (2025).","journal-title":"Nature"},{"key":"76210_CR48","doi-asserted-by":"publisher","first-page":"444","DOI":"10.1105\/tpc.111.092791","volume":"24","author":"S Torti","year":"2012","unstructured":"Torti, S. et al. Analysis of the Arabidopsis shoot meristem transcriptome during floral transition identifies distinct regulatory patterns and a leucine-rich repeat protein that promotes flowering. Plant Cell. 24, 444\u2013462 (2012).","journal-title":"Plant Cell."},{"key":"76210_CR49","doi-asserted-by":"publisher","first-page":"738","DOI":"10.1016\/j.cell.2009.06.014","volume":"138","author":"JW Wang","year":"2009","unstructured":"Wang, J. W., Czech, B. & Weigel, D. miR156-regulated SPL transcription factors define an endogenous flowering pathway in Arabidopsis thaliana. Cell 138, 738\u2013749 (2009).","journal-title":"Cell"},{"key":"76210_CR50","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1104\/pp.16.01523","volume":"173","author":"Y Hyun","year":"2017","unstructured":"Hyun, Y., Richter, R. & Coupland, G. Competence to flower: Age-controlled sensitivity to environmental cues. Plant Physiol. 173, 36\u201346 (2017).","journal-title":"Plant Physiol."},{"key":"76210_CR51","doi-asserted-by":"publisher","first-page":"683","DOI":"10.1111\/j.1365-313X.2006.02983.x","volume":"49","author":"M Gandikota","year":"2007","unstructured":"Gandikota, M. et al. The miRNA156\/157 recognition element in the 3\u2032 UTR of the Arabidopsis SBP box gene SPL3 prevents early flowering by translational inhibition in seedlings. Plant J. 49, 683\u2013693 (2007).","journal-title":"Plant J."},{"key":"76210_CR52","doi-asserted-by":"publisher","first-page":"254","DOI":"10.1016\/j.devcel.2016.04.001","volume":"37","author":"Y Hyun","year":"2016","unstructured":"Hyun, Y. et al. Multi-layered regulation of SPL15 and cooperation with SOC1 integrate endogenous flowering pathways at the Arabidopsis shoot meristem. Dev. Cell 37, 254\u2013266 (2016).","journal-title":"Dev. Cell"},{"key":"76210_CR53","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1016\/j.devcel.2009.06.007","volume":"17","author":"A Yamaguchi","year":"2009","unstructured":"Yamaguchi, A. et al. The microRNA-regulated SBP-box transcription factor SPL3 is a direct upstream activator of LEAFY, FRUITFULL, and APETALA1. Dev. Cell 17, 268\u2013278 (2009).","journal-title":"Dev. Cell"},{"key":"76210_CR54","doi-asserted-by":"publisher","first-page":"2022","DOI":"10.1126\/science.1088060","volume":"303","author":"X Chen","year":"2004","unstructured":"Chen, X. A microRNA as a translational repressor of APETALA2 in Arabidopsis flower development. Science 303, 2022\u20132025 (2004).","journal-title":"Science"},{"key":"76210_CR55","doi-asserted-by":"publisher","first-page":"e1000148","DOI":"10.1371\/journal.pbio.1000148","volume":"7","author":"J Mathieu","year":"2009","unstructured":"Mathieu, J., Yant, L. J., M\u00fcrdter, F., K\u00fcttner, F. & Schmid, M. Repression of flowering by the miR172 target SMZ. PLoS Biol. 7, e1000148 (2009).","journal-title":"PLoS Biol."},{"key":"76210_CR56","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-03067-5","volume":"9","author":"V Balanz\u00e0","year":"2018","unstructured":"Balanz\u00e0, V. et al. Genetic control of meristem arrest and life span in Arabidopsis by a FRUITFULL-APETALA2 pathway. Nat. Commun. 9, 565 (2018).","journal-title":"Nat. Commun."},{"key":"76210_CR57","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1111\/nph.18111","volume":"235","author":"Q Sang","year":"2022","unstructured":"Sang, Q. et al. MicroRNA172 controls inflorescence meristem size through regulation of APETALA2 in Arabidopsis. New Phytol. 235, 356\u2013371 (2022).","journal-title":"New Phytol."},{"key":"76210_CR58","doi-asserted-by":"publisher","first-page":"820","DOI":"10.1105\/tpc.113.109355","volume":"25","author":"KE Jaeger","year":"2013","unstructured":"Jaeger, K. E., Pullen, N., Lamzin, S., Morris, R. J. & Wigge, P. A. Interlocking feedback loops govern the dynamic behavior of the floral transition in Arabidopsis. Plant Cell 25, 820\u2013833 (2013).","journal-title":"Plant Cell"},{"key":"76210_CR59","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1242\/dev.127.4.725","volume":"127","author":"C Ferr\u00e1ndiz","year":"2000","unstructured":"Ferr\u00e1ndiz, C., Gu, Q., Martienssen, R. & Yanofsky, M. F. Redundant regulation of meristem identity and plant architecture by FRUITFULL, APETALA1 and CAULIFLOWER. Development 127, 725\u2013734 (2000).","journal-title":"Development"},{"key":"76210_CR60","doi-asserted-by":"publisher","first-page":"1489","DOI":"10.1038\/ng.253","volume":"40","author":"S Melzer","year":"2008","unstructured":"Melzer, S. et al. Flowering-time genes modulate meristem determinacy and growth form in Arabidopsis thaliana. Nat. Genet. 40, 1489\u20131492 (2008).","journal-title":"Nat. Genet."},{"key":"76210_CR61","doi-asserted-by":"publisher","first-page":"E2760","DOI":"10.1073\/pnas.1409567111","volume":"111","author":"F Andr\u00e9s","year":"2014","unstructured":"Andr\u00e9s, F. et al. SHORT VEGETATIVE PHASE reduces gibberellin biosynthesis at the Arabidopsis shoot apex to regulate the floral transition. Proc. Natl. Acad. Sci. USA. 111, E2760\u2013E2769 (2014).","journal-title":"Proc. Natl. Acad. Sci. USA."},{"key":"76210_CR62","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1016\/0065-2571(65)90067-1","volume":"3","author":"BC Goodwin","year":"1965","unstructured":"Goodwin, B. C. Oscillatory behavior in enzymatic control processes. Adv. Enzyme Regul. 3, 425\u2013438 (1965).","journal-title":"Adv. Enzyme Regul."},{"key":"76210_CR63","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1038\/35002125","volume":"403","author":"MB Elowitz","year":"2000","unstructured":"Elowitz, M. B. & Leibler, S. A synthetic oscillatory network of transcriptional regulators. Nature 403, 335\u2013338 (2000).","journal-title":"Nature"},{"key":"76210_CR64","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1016\/j.cels.2018.02.008","volume":"6","author":"R Perez-Carrasco","year":"2018","unstructured":"Perez-Carrasco, R. et al. Combining a toggle switch and a repressilator within the AC-DC circuit generates distinct dynamical behaviors. Cell Syst. 6, 521\u2013530.e3 (2018).","journal-title":"Cell Syst."},{"key":"76210_CR65","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1016\/j.devcel.2005.01.018","volume":"8","author":"R Schwab","year":"2005","unstructured":"Schwab, R. et al. Specific effects of microRNAs on the plant transcriptome. Dev. Cell 8, 517\u2013527 (2005).","journal-title":"Dev. Cell"},{"key":"76210_CR66","doi-asserted-by":"publisher","first-page":"2730","DOI":"10.1105\/tpc.016238","volume":"15","author":"MJ Aukerman","year":"2003","unstructured":"Aukerman, M. J. & Sakai, H. Regulation of flowering time and floral organ identity by a microRNA and its APETALA2-like target genes. Plant Cell 15, 2730\u20132741 (2003).","journal-title":"Plant Cell"},{"key":"76210_CR67","doi-asserted-by":"publisher","DOI":"10.1103\/PhysRevE.111.024213","volume":"111","author":"M Fucho-Rius","year":"2025","unstructured":"Fucho-Rius, M. et al. Local nearby bifurcations lead to synergies in critical slowing down: the case of mushroom bifurcations. Phys. Rev. E. 111, 024213 (2025).","journal-title":"Phys. Rev. E."},{"key":"76210_CR68","doi-asserted-by":"publisher","first-page":"e2000640","DOI":"10.1371\/journal.pbio.2000640","volume":"14","author":"M Mojtahedi","year":"2016","unstructured":"Mojtahedi, M. et al. Cell fate decision as high-dimensional critical state transition. PLoS Biol 14, e2000640 (2016).","journal-title":"PLoS Biol"},{"key":"76210_CR69","doi-asserted-by":"publisher","first-page":"10168","DOI":"10.1073\/pnas.1817853116","volume":"116","author":"JP Fouracre","year":"2019","unstructured":"Fouracre, J. P. & Poethig, R. S. Role for the shoot apical meristem in the specification of juvenile leaf identity in Arabidopsis. Proc. Natl. Acad. Sci. USA. 116, 10168\u201310177 (2019).","journal-title":"Proc. Natl. Acad. Sci. USA."},{"key":"76210_CR70","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1038\/nrg2102","volume":"8","author":"U Alon","year":"2007","unstructured":"Alon, U. Network motifs: theory and experimental approaches. Nat. Rev. Genet. 8, 450\u2013461 (2007).","journal-title":"Nat. Rev. Genet."},{"key":"76210_CR71","doi-asserted-by":"publisher","first-page":"e1005154","DOI":"10.1371\/journal.pcbi.1005154","volume":"12","author":"R Perez-Carrasco","year":"2016","unstructured":"Perez-Carrasco, R., Guerrero, P., Briscoe, J. & Page, K. M. Intrinsic noise profoundly alters the dynamics and steady state of morphogen-controlled bistable genetic switches. PLoS Comput. Biol. 12, e1005154 (2016).","journal-title":"PLoS Comput. Biol."},{"key":"76210_CR72","doi-asserted-by":"publisher","unstructured":"Maretvadakethope, S. et al. Let there be multifunctionality: uncovering the criticality zoo of the AC-DC genetic circuit. Preprint at https:\/\/doi.org\/10.1101\/2025.08.22.671785 (2025).","DOI":"10.1101\/2025.08.22.671785"},{"key":"76210_CR73","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/j.cell.2011.10.047","volume":"148","author":"N Balaskas","year":"2012","unstructured":"Balaskas, N. et al. Gene regulatory logic for reading the Sonic Hedgehog signaling gradient in the vertebrate neural tube. Cell 148, 273\u2013284 (2012).","journal-title":"Cell"},{"key":"76210_CR74","doi-asserted-by":"publisher","first-page":"e2003174","DOI":"10.1371\/journal.pbio.2003174","volume":"16","author":"B Verd","year":"2018","unstructured":"Verd, B. et al. A damped oscillator imposes temporal order on posterior gap gene expression in Drosophila. PLoS Biol. 16, e2003174 (2018).","journal-title":"PLoS Biol."},{"key":"76210_CR75","doi-asserted-by":"publisher","first-page":"e1007969","DOI":"10.1371\/journal.pcbi.1007969","volume":"16","author":"S Nagata","year":"2020","unstructured":"Nagata, S. & Kikuchi, M. Emergence of cooperative bistability and robustness of gene regulatory networks. PLoS Comput. Biol. 16, e1007969 (2020).","journal-title":"PLoS Comput. Biol."},{"key":"76210_CR76","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.59485","volume":"10","author":"K Abley","year":"2021","unstructured":"Abley, K. et al. An ABA-GA bistable switch can account for natural variation in the variability of Arabidopsis seed germination time. eLife 10, e59485 (2021).","journal-title":"eLife"},{"key":"76210_CR77","doi-asserted-by":"publisher","first-page":"20230048","DOI":"10.1098\/rstb.2023.0048","volume":"379","author":"K Abley","year":"2024","unstructured":"Abley, K., Goswami, R. & Locke, J. C. W. Bet-hedging and variability in plant development: seed germination and beyond. Philos. Trans. R. Soc. B. Biol. Sci. 379, 20230048 (2024).","journal-title":"Philos. Trans. R. Soc. B. Biol. Sci."},{"key":"76210_CR78","doi-asserted-by":"publisher","first-page":"2587","DOI":"10.1093\/jxb\/eri254","volume":"56","author":"F Tooke","year":"2005","unstructured":"Tooke, F., Ordidge, M., Chiurugwi, T. & Battey, N. Mechanisms and function of flower and inflorescence reversion. J. Exp. Bot. 56, 2587\u20132599 (2005).","journal-title":"J. Exp. Bot."},{"key":"76210_CR79","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1111\/j.1469-8137.2009.03141.x","volume":"186","author":"E McCullough","year":"2010","unstructured":"McCullough, E. et al. Photoperiod-dependent floral reversion in the natural allopolyploid Arabidopsis suecica. New Phytol 186, 239\u2013250 (2010).","journal-title":"New Phytol"},{"key":"76210_CR80","doi-asserted-by":"publisher","first-page":"e1005285","DOI":"10.1371\/journal.pcbi.1005285","volume":"13","author":"B Verd","year":"2017","unstructured":"Verd, B., Crombach, A. & Jaeger, J. Dynamic maternal gradients control timing and shift-rates for Drosophila Gap gene expression. PLoS Comput. Biol. 13, e1005285 (2017).","journal-title":"PLoS Comput. Biol."},{"key":"76210_CR81","doi-asserted-by":"publisher","first-page":"1193","DOI":"10.1093\/jxb\/ert482","volume":"65","author":"V Balanz\u00e0","year":"2014","unstructured":"Balanz\u00e0, V., Mart\u00ednez-Fern\u00e1ndez, I. & Ferr\u00e1ndiz, C. Sequential action of FRUITFULL as a modulator of the activity of the floral regulators SVP and SOC1. J. Exp. Bot. 65, 1193\u20131203 (2014).","journal-title":"J. Exp. Bot."},{"key":"76210_CR82","doi-asserted-by":"publisher","first-page":"150069","DOI":"10.1016\/j.bbrc.2024.150069","volume":"720","author":"D Koch","year":"2024","unstructured":"Koch, D., Nandan, A., Ramesan, G. & Koseska, A. Biological computations: Limitations of attractor-based formalisms and the need for transients. Biochem. Biophys. Res. Commun. 720, 150069 (2024).","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"76210_CR83","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1016\/0022-5193(68)90189-6","volume":"20","author":"JS Griffith","year":"1968","unstructured":"Griffith, J. S. Mathematics of cellular control processes I. Negative feedback to one gene. J. Theor. Biol. 20, 202\u2013208 (1968).","journal-title":"J. Theor. Biol."},{"key":"76210_CR84","unstructured":"Ingalls, B. P. Mathematical Modelling in Systems Biology: An Introduction. (MIT press, 2013)."},{"key":"76210_CR85","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1109\/TASSP.1978.1163055","volume":"26","author":"H Sakoe","year":"1978","unstructured":"Sakoe, H. & Chiba, S. Dynamic programming algorithm optimization for spoken word recognition. IEEE Trans. Acoust. Speech Signal Process. 26, 43\u201349 (1978).","journal-title":"IEEE Trans. Acoust. Speech Signal Process."},{"key":"76210_CR86","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1016\/0165-1684(82)90009-3","volume":"4","author":"KK Paliwal","year":"1982","unstructured":"Paliwal, K. K., Agarwal, A. & Sinha, S. S. A modification over Sakoe and Chiba\u2019s dynamic time warping algorithm for isolated word recognition. Signal Process 4, 329\u2013333 (1982).","journal-title":"Signal Process"},{"key":"76210_CR87","unstructured":"Clewley, R., Sherwood, W. E., LaMar, M. & Guckenheimer, J. PyDSTool, a software environment for dynamical systems modeling. (2007)."},{"key":"76210_CR88","doi-asserted-by":"crossref","unstructured":"Gardiner, C. W. Handbook of Stochastic Methods for Physics, Chemistry and the Natural Sciences. (Springer Berlin Heidelberg, 1985).","DOI":"10.1007\/978-3-662-02452-2"},{"key":"76210_CR89","doi-asserted-by":"publisher","first-page":"2366","DOI":"10.1101\/gad.813600","volume":"14","author":"H Lee","year":"2000","unstructured":"Lee, H. et al. The AGAMOUS-LIKE 20 MADS domain protein integrates floral inductive pathways in Arabidopsis. Genes Dev. 14, 2366\u20132376 (2000).","journal-title":"Genes Dev."},{"key":"76210_CR90","first-page":"4168","volume":"142","author":"D Kurihara","year":"2015","unstructured":"Kurihara, D., Mizuta, Y., Sato, Y. & Higashiyama, T. ClearSee: a rapid optical clearing reagent for whole-plant fluorescence imaging. Development 142, 4168\u20134179 (2015).","journal-title":"Development"},{"key":"76210_CR91","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1007\/s00497-015-0267-1","volume":"28","author":"TJ Musielak","year":"2015","unstructured":"Musielak, T. J., Schenkel, L., Kolb, M., Henschen, A. & Bayer, M. A simple and versatile cell wall staining protocol to study plant reproduction. Plant Reprod 28, 161\u2013169 (2015).","journal-title":"Plant Reprod"},{"key":"76210_CR92","doi-asserted-by":"crossref","unstructured":"Weigert, M., Schmidt, U., Haase, R., Sugawara, K. & Myers, G. Star-convex polyhedra for 3D object detection and segmentation in microscopy. in 2020 IEEE Winter Conference on Applications of Computer Vision (WACV), 3655\u20133662 (2020).","DOI":"10.1109\/WACV45572.2020.9093435"},{"key":"76210_CR93","doi-asserted-by":"publisher","DOI":"10.1242\/dev.202800","volume":"151","author":"A Vijayan","year":"2024","unstructured":"Vijayan, A. et al. A deep learning-based toolkit for 3D nuclei segmentation and quantitative analysis in cellular and tissue context. Development 151, dev202800 (2024).","journal-title":"Development"},{"key":"76210_CR94","doi-asserted-by":"publisher","unstructured":"Yu, Q. StarDist Plant Nuclei 3D ResNet. Zenodo https:\/\/doi.org\/10.5281\/zenodo.8432366 (2023).","DOI":"10.5281\/zenodo.8432366"}],"container-title":["Nature Communications"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-76210-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-76210-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-76210-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T14:59:13Z","timestamp":1787669953000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-76210-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,25]]},"references-count":94,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["76210"],"URL":"https:\/\/doi.org\/10.1038\/s41467-026-76210-2","relation":{},"ISSN":["2041-1723"],"issn-type":[{"value":"2041-1723","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,25]]},"assertion":[{"value":"25 November 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 July 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 August 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}}],"article-number":"8973"}}