{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,22]],"date-time":"2026-07-22T23:34:42Z","timestamp":1784763282158,"version":"3.55.0"},"reference-count":90,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,1,5]],"date-time":"2026-01-05T00:00:00Z","timestamp":1767571200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,1,5]],"date-time":"2026-01-05T00:00:00Z","timestamp":1767571200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-025-29571-5","type":"journal-article","created":{"date-parts":[[2026,1,5]],"date-time":"2026-01-05T13:57:03Z","timestamp":1767621423000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["The sexual reproductive strategy in clonal Scopolia carniolica is based on obligatory cross-pollination and self-incompatibility"],"prefix":"10.1038","volume":"16","author":[{"given":"Karolina","family":"Tymoszuk-Rydzewska","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bo\u017cena","family":"Denisow","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jacek","family":"Mielniczuk","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ernest","family":"Stawiarz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,1,5]]},"reference":[{"key":"29571_CR1","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/j.cois.2015.11.002","volume":"12","author":"D Senapathi","year":"2015","unstructured":"Senapathi, D. et al. Pollinator conservation\u2014the difference between managing for pollination services and preserving pollinator diversity. Curr. Opin. Insect Sci. 12, 93\u2013101 (2015).","journal-title":"Curr. Opin. Insect Sci."},{"key":"29571_CR2","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1111\/j.1438-8677.2011.00510.x","volume":"14","author":"M Zych","year":"2012","unstructured":"Zych, M. & Stpiczy\u0144ska, M. Neither protogynous nor obligatory out-crossed: pollination biology and breeding system of the European red list Fritillaria meleagris L. (Liliaceae). Plant. Biol. 14, 285\u2013294 (2012).","journal-title":"Plant. Biol."},{"key":"29571_CR3","doi-asserted-by":"publisher","first-page":"1231","DOI":"10.1007\/s00606-018-1543-y","volume":"304","author":"S Anto\u0144","year":"2018","unstructured":"Anto\u0144, S. & Denisow, B. Pollination biology and breeding system in five nocturnal species of oenothera (Onagraceae): reproductive assurance and opportunities for outcrossing. Plant. Syst. Evol. 304, 1231\u20131243 (2018).","journal-title":"Plant. Syst. Evol."},{"key":"29571_CR4","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1098\/rstb.2009.0199","volume":"365","author":"SCH Barrett","year":"2010","unstructured":"Barrett, S. C. H. Understanding plant reproductive diversity. Philos. Trans. R Soc. B Biol. Sci. 365, 99\u2013109 (2010).","journal-title":"Philos. Trans. R Soc. B Biol. Sci."},{"key":"29571_CR5","doi-asserted-by":"publisher","first-page":"338","DOI":"10.1038\/s41437-023-00649-y","volume":"131","author":"Z Diaz-Martin","year":"2023","unstructured":"Diaz-Martin, Z. et al. Reproductive strategies and their consequences for divergence, gene flow, and genetic diversity in three taxa of Clarkia. Heredity 131, 338\u2013349 (2023).","journal-title":"Heredity"},{"key":"29571_CR6","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1111\/j.1600-0706.2010.18644.x","volume":"120","author":"J Ollerton","year":"2011","unstructured":"Ollerton, J., Winfree, R. & Tarrant, S. How many flowering plants are pollinated by animals? Oikos 120, 321\u2013326 (2011).","journal-title":"Oikos"},{"key":"29571_CR7","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1111\/j.1558-5646.1977.tb00979.x","volume":"31","author":"RW Cruden","year":"1977","unstructured":"Cruden, R. W. Pollen-Ovule ratios: A Conservative indicator of breeding systems in flowering plants. Evolution 31, 32 (1977).","journal-title":"Evolution"},{"key":"29571_CR8","doi-asserted-by":"publisher","first-page":"7037","DOI":"10.1073\/pnas.97.13.7037","volume":"97","author":"EK Holsinger","year":"2000","unstructured":"Holsinger, E. K Reproductive systems and evolution in vascular plants. Proc. Natl. Acad. Sci. 97, 7037\u20137042 (2000).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"29571_CR9","doi-asserted-by":"publisher","first-page":"991","DOI":"10.1098\/rstb.2003.1301","volume":"358","author":"SCH Barrett","year":"2003","unstructured":"Barrett, S. C. H. Mating strategies in flowering plants: the outcrossing\u2013selfing paradigm and beyond. Philos. Trans. R Soc. Lond. B Biol. Sci. 358, 991\u20131004 (2003).","journal-title":"Philos. Trans. R Soc. Lond. B Biol. Sci."},{"key":"29571_CR10","doi-asserted-by":"publisher","first-page":"717","DOI":"10.1111\/nph.12874","volume":"203","author":"PE Gibbs","year":"2014","unstructured":"Gibbs, P. E. Late-acting self\u2010incompatibility \u2013 the pariah breeding system in flowering plants. New. Phytol. 203, 717\u2013734 (2014).","journal-title":"New. Phytol"},{"key":"29571_CR11","doi-asserted-by":"crossref","unstructured":"Willmer, P. Pollination and Floral Ecology (Princeton University Press, 2011).","DOI":"10.23943\/princeton\/9780691128610.001.0001"},{"key":"29571_CR12","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1146\/annurev.ecolsys.36.091704.175539","volume":"36","author":"C Goodwillie","year":"2005","unstructured":"Goodwillie, C., Kalisz, S. & Eckert, C. G. The evolutionary enigma of mixed mating systems in plants: Occurrence, theoretical Explanations, and empirical evidence. Annu. Rev. Ecol. Evol. Syst. 36, 47\u201379 (2005).","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"29571_CR13","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1007\/BF02856594","volume":"60","author":"M Burd","year":"1994","unstructured":"Burd, M. Bateman\u2019s principle and plant reproduction: the role of pollen limitation in fruit and seed set. Bot. Rev. 60, 83\u2013139 (1994).","journal-title":"Bot. Rev."},{"key":"29571_CR14","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1111\/j.1095-8312.2000.tb01221.x","volume":"69","author":"BMH Larson","year":"2000","unstructured":"Larson, B. M. H. & Barrett, S. C. H. A comparative analysis of pollen limitation in flowering plants. Biol. J. Linn. Soc. 69, 503\u2013520 (2000).","journal-title":"Biol. J. Linn. Soc."},{"key":"29571_CR15","doi-asserted-by":"publisher","first-page":"1924","DOI":"10.1111\/j.0014-3820.2004.tb00480.x","volume":"58","author":"M Vallejo-Mar\u00edn","year":"2004","unstructured":"Vallejo-Mar\u00edn, M. & Uyenoyama, M. K. On the evolutionary costs of self-incompatibility: incomplete reproductive compensation due to pollen limitation. Evolution 58, 1924\u20131935 (2004).","journal-title":"Evolution"},{"key":"29571_CR16","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1146\/annurev.ecolsys.36.102403.115320","volume":"36","author":"TM Knight","year":"2005","unstructured":"Knight, T. M. et al. Pollen limitation of plant reproduction: pattern and process. Annu. Rev. Ecol. Evol. Syst. 36, 467\u2013497 (2005).","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"29571_CR17","doi-asserted-by":"publisher","first-page":"1641","DOI":"10.1111\/j.1558-5646.1991.tb02669.x","volume":"45","author":"DH Les","year":"1991","unstructured":"Les, D. H., Reinartz, J. A. & Esselman, E. J. Genetic consequences of rarity in Aster furcatus (asteraceae), a threatened, self-incompatible plant. Evolution 45, 1641\u20131650 (1991).","journal-title":"Evolution"},{"key":"29571_CR18","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1146\/annurev.arplant.56.032604.144249","volume":"56","author":"S Takayama","year":"2005","unstructured":"Takayama, S. & Isogai, A. Self-incompatibility in plants. Annu. Rev. Plant. Biol. 56, 467\u2013489 (2005).","journal-title":"Annu. Rev. Plant. Biol."},{"key":"29571_CR19","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/ery406","author":"L Wang","year":"2018","unstructured":"Wang, L. et al. Self-incompatibility in Papaver pollen: programmed cell death in an acidic environment. J. Exp. Bot. https:\/\/doi.org\/10.1093\/jxb\/ery406 (2018).","journal-title":"J. Exp. Bot."},{"key":"29571_CR20","doi-asserted-by":"publisher","first-page":"264","DOI":"10.2183\/pjab.100.014","volume":"100","author":"K Murase","year":"2024","unstructured":"Murase, K., Takayama, S. & Isogai, A. Molecular mechanisms of self-incompatibility in Brassicaceae and Solanaceae. Proc. Jpn Acad. Ser. B 100, 264\u2013280 (2024).","journal-title":"Proc. Jpn Acad. Ser. B"},{"key":"29571_CR21","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1007\/BF00985456","volume":"225","author":"JL Vesprini","year":"2000","unstructured":"Vesprini, J. L. & Galetto, L. The reproductive biology OfJaborosa integrifolia (Solanaceae): why its fruits are so rare? Plant. Syst. Evol. 225, 15\u201328 (2000).","journal-title":"Plant. Syst. Evol."},{"key":"29571_CR22","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1086\/523362","volume":"169","author":"B Igic","year":"2008","unstructured":"Igic, B., Lande, R. & Kohn, J. R. Loss of Self-Incompatibility and its evolutionary consequences. Int. J. Plant. Sci. 169, 93\u2013104 (2008).","journal-title":"Int. J. Plant. Sci."},{"key":"29571_CR23","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1017\/S0266467411000289","volume":"27","author":"MA Caraballo-Ortiz","year":"2011","unstructured":"Caraballo-Ortiz, M. A., Santiago-Valent\u00edn, E. & Carlo, T. A. Flower number and distance to neighbours affect the fecundity of Goetzea elegans (Solanaceae). J. Trop. Ecol. 27, 521\u2013528 (2011).","journal-title":"J. Trop. Ecol."},{"key":"29571_CR24","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1007\/BF02861001","volume":"52","author":"SR Seavey","year":"1986","unstructured":"Seavey, S. R. & Bawa, K. S. Late-acting self-incompatibility in angiosperms. Bot. Rev. 52, 195\u2013219 (1986).","journal-title":"Bot. Rev."},{"key":"29571_CR25","doi-asserted-by":"publisher","first-page":"415","DOI":"10.1111\/j.1469-8137.2006.01924.x","volume":"173","author":"M Vallejo-Mar\u00edn","year":"2007","unstructured":"Vallejo-Mar\u00edn, M. & O\u2019Brien, H. E. Correlated evolution of self\u2010incompatibility and clonal reproduction in Solanum (Solanaceae). New. Phytol. 173, 415\u2013421 (2007).","journal-title":"New. Phytol"},{"key":"29571_CR26","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1006\/anbo.1999.1033","volume":"85","author":"A Stephenson","year":"2000","unstructured":"Stephenson, A. Interrelationships among inbreeding Depression, plasticity in the Self-incompatibility system, and the breeding system of Campanula rapunculoides L. (Campanulaceae). Ann. Bot. 85, 211\u2013219 (2000).","journal-title":"Ann. Bot."},{"key":"29571_CR27","doi-asserted-by":"publisher","first-page":"e16150","DOI":"10.1002\/ajb2.16150","volume":"110","author":"C Pretz","year":"2023","unstructured":"Pretz, C. & Smith, S. D. How does a self-incompatible individual transition to self\u2010compatibility during its lifetime? Am. J. Bot. 110, e16150 (2023).","journal-title":"Am. J. Bot."},{"key":"29571_CR28","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1007\/s00497-024-00517-7","volume":"38","author":"C Pretz","year":"2025","unstructured":"Pretz, C., Au, E. & Smith, S. D. Decay of self-incompatibility within a lifespan in physalis acutifolia (Solanaceae). Plant. Reprod. 38, 7 (2025).","journal-title":"Plant. Reprod."},{"key":"29571_CR29","doi-asserted-by":"publisher","first-page":"46","DOI":"10.26786\/1920-7603(2013)5","volume":"11","author":"R Lundgren","year":"2013","unstructured":"Lundgren, R., Laz\u00e1ro, A. & Totland, \u00d8. Experimental pollinator decline affects plant reproduction and is mediated by plant mating system. J. Pollinat. Ecol. 11, 46\u201356 (2013).","journal-title":"J. Pollinat. Ecol."},{"key":"29571_CR30","doi-asserted-by":"publisher","first-page":"296","DOI":"10.1038\/s41559-024-02595-2","volume":"9","author":"M Artamendi","year":"2024","unstructured":"Artamendi, M., Martin, P. A., Bartomeus, I. & Magrach, A. Loss of pollinator diversity consistently reduces reproductive success for wild and cultivated plants. Nat. Ecol. Evol. 9, 296\u2013313 (2024).","journal-title":"Nat. Ecol. Evol."},{"key":"29571_CR31","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1146\/annurev.ecolsys.29.1.83","volume":"29","author":"CA Kearns","year":"1998","unstructured":"Kearns, C. A., Inouye, D. W. & Waser, N. M. Endangered mutualisms: the conservation of Plant-Pollinator interactions. Annu. Rev. Ecol. Syst. 29, 83\u2013112 (1998).","journal-title":"Annu. Rev. Ecol. Syst."},{"key":"29571_CR32","unstructured":"Plants of the World Online | Kew Science. Plants of the World Online. https:\/\/powo.science.kew.org\/results?q=Scopolia%20carniolica"},{"key":"29571_CR33","unstructured":"Group), S. K. & (Cave, I. S. IUCN red list of threatened species: Scopolia carniolica. IUCN Red List. Threat Species (2013). https:\/\/www.iucnredlist.org\/en"},{"key":"29571_CR34","unstructured":"Dziennik Ustaw. poz. (2014). 1409.\u00a0https:\/\/dziennikustaw.gov.pl\/du\/2014\/1409"},{"key":"29571_CR35","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1007\/s11829-024-10059-4","volume":"18","author":"K Tymoszuk","year":"2024","unstructured":"Tymoszuk, K., Dmitruk, M., Jachu\u0142a, J. & Denisow, B. Flower structure and floral reward in scopolia carniolica (Solanaceae) \u2013 is it a plant that can support the bumblebee food base in early spring? Arthropod-Plant Interact. 18, 403\u2013416 (2024).","journal-title":"Arthropod-Plant Interact."},{"key":"29571_CR36","unstructured":"Flora Europaea. vol. 3: Diapensiaceae to Myoporaceae (ed. Tutin, T. G.) (Cambridge University Press, Cambridge, 2010)."},{"key":"29571_CR37","doi-asserted-by":"publisher","first-page":"262","DOI":"10.2307\/2394758","volume":"59","author":"WG D\u2019Arcy","year":"1972","unstructured":"D\u2019Arcy, W. G. & Solanaceae Studies, I. I. Typification of subdivisions of solanum. Ann. Mo Bot. Gard. 59, 262 (1972).","journal-title":"Ann. Mo Bot. Gard"},{"key":"29571_CR38","doi-asserted-by":"publisher","first-page":"124","DOI":"10.2307\/3391672","volume":"2","author":"WG D\u2019Arcy","year":"1992","unstructured":"D\u2019Arcy, W. G. Zhi-yun, Z. Notes on the Solanaceae of China and neighboring areas. Novon 2, 124 (1992).","journal-title":"Novon"},{"key":"29571_CR39","unstructured":"Knuth, P. Handbuch Der Bl\u00fctenbiologie. (Engelmann, 1899)."},{"key":"29571_CR40","doi-asserted-by":"publisher","first-page":"93","DOI":"10.5586\/asbp.1991.007","volume":"60","author":"A Shehata","year":"2014","unstructured":"Shehata, A. & Zenkteler, M. In vitro placental self and cross pollination in some species. Acta Soc. Bot. Pol. 60, 93\u201397 (2014).","journal-title":"Acta Soc. Bot. Pol."},{"key":"29571_CR41","doi-asserted-by":"publisher","first-page":"225","DOI":"10.17951\/pjss.2019.52.2.225","volume":"52","author":"K Wo\u017anica","year":"2019","unstructured":"Wo\u017anica, K., J\u00f3zefowska, A., Soko\u0142owska, J., Mazurek, R. & Zaleski, T. Classification of brown earths based on field and laboratory properties: problematic issues and proposition of their solution. Pol. J. Soil. Sci. 52, 225 (2019).","journal-title":"Pol. J. Soil. Sci."},{"key":"29571_CR42","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1127\/0941-2948\/2011\/0285","volume":"20","author":"F Rubel","year":"2011","unstructured":"Rubel, F. & Kottek, M. Comments on: \u2018The thermal zones of the earth\u2019 by Wladimir K\u00f6ppen (1884). Meteorol. Z. 20, 361\u2013365 (2011).","journal-title":"Meteorol. Z."},{"key":"29571_CR43","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1007\/s004970050138","volume":"11","author":"A Dafni","year":"1998","unstructured":"Dafni, A. & Mau\u00e9s, M. M. A rapid and simple procedure to determine stigma receptivity. Sex. Plant. Reprod. 11, 177\u2013180 (1998).","journal-title":"Sex. Plant. Reprod."},{"key":"29571_CR44","unstructured":"Laboratory Techniques in Cellular and Molecular Medicine ((Palack\u00fd University Olomouc, Olomouc, 2021)."},{"key":"29571_CR45","doi-asserted-by":"publisher","first-page":"139","DOI":"10.5586\/aa.2006.014","volume":"59","author":"B Denisow","year":"2012","unstructured":"Denisow, B. & Bo\u017cek, M. Blooming biology and pollen abundance of Anemone japonica Houtt.= anemone x hybrida hort. Acta Agrobot 59, 139\u2013146 (2012).","journal-title":"Acta Agrobot"},{"key":"29571_CR46","unstructured":"Analyzeseeds \u2013 AOSA | SCST, Tetrazolium Testing Handbook. https:\/\/analyzeseeds.com\/"},{"key":"29571_CR47","doi-asserted-by":"crossref","unstructured":"Chapter 7: Seed health testing. Int. Rules Seed Test. 2024, 1\u201314 (2024).","DOI":"10.15258\/istarules.2023.07"},{"key":"29571_CR48","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1007\/s00606-006-0493-y","volume":"263","author":"M Zych","year":"2007","unstructured":"Zych, M. On flower visitors and true pollinators: the case of protandrous heracleum sphondylium L. (Apiaceae). Plant. Syst. Evol. 263, 159\u2013179 (2007).","journal-title":"Plant. Syst. Evol."},{"key":"29571_CR49","unstructured":"Dafni, A. Pollination Ecology: A Practical Approach. (IRL Press at Oxford University Press, 1992)."},{"key":"29571_CR50","doi-asserted-by":"publisher","unstructured":"Venables, W. N. & Ripley, B. D. Modern Applied Statistics with S (Springer New York, 2002). https:\/\/doi.org\/10.1007\/978-0-387-21706-2","DOI":"10.1007\/978-0-387-21706-2"},{"key":"29571_CR51","doi-asserted-by":"crossref","unstructured":"Boucher, J. J., Ireland, H. S., Wang, R., David, K. M. & Schaffer, R. J. The genetic control of herkogamy. Funct Plant. Biol 51\u00a0(2024).","DOI":"10.1071\/FP23315"},{"key":"29571_CR52","doi-asserted-by":"publisher","first-page":"643","DOI":"10.1126\/science.1148428","volume":"318","author":"KY Chen","year":"2007","unstructured":"Chen, K. Y., Cong, B., Wing, R., Vrebalov, J. & Tanksley, S. D. Changes in regulation of a transcription factor lead to autogamy in cultivated tomatoes. Science 318, 643\u2013645 (2007).","journal-title":"Science"},{"key":"29571_CR53","doi-asserted-by":"publisher","first-page":"358","DOI":"10.1086\/297040","volume":"153","author":"DG Lloyd","year":"1992","unstructured":"Lloyd, D. G. & Schoen, D. J. Self- and Cross-Fertilization in Plants. I. Functional dimensions. Int. J. Plant. Sci. 153, 358\u2013369 (1992).","journal-title":"Int. J. Plant. Sci."},{"key":"29571_CR54","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1080\/0028825X.1986.10409725","volume":"24","author":"DG Lloyd","year":"1986","unstructured":"Lloyd, D. G. & Webb, C. J. The avoidance of interference between the presentation of pollen and stigmas in angiosperms I. Dichogamy. N Z. J. Bot. 24, 135\u2013162 (1986).","journal-title":"N Z. J. Bot."},{"key":"29571_CR55","first-page":"557","volume":"80","author":"RI Bertin","year":"1993","unstructured":"Bertin, R. I. Incidence of monoecy and dichogamy in relation to self-fertilization in angiosperms. Am. J. Bot. 80, 557\u2013560 (1993).","journal-title":"Am. J. Bot."},{"key":"29571_CR56","doi-asserted-by":"publisher","first-page":"367","DOI":"10.1007\/BF00014943","volume":"22","author":"AJ De Jong","year":"1993","unstructured":"De Jong, A. J., Schmidt, E. D. L. & De Vries, S. C. Early events in higher-plant embryogenesis. Plant. Mol. Biol. 22, 367\u2013377 (1993).","journal-title":"Plant. Mol. Biol."},{"key":"29571_CR57","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1046\/j.1365-2540.1998.00415.x","volume":"81","author":"D Charlesworth","year":"1998","unstructured":"Charlesworth, D. & Yang, Z. Allozyme diversity in leavenworthia populations with different inbreeding levels. Heredity 81, 453\u2013461 (1998).","journal-title":"Heredity"},{"key":"29571_CR58","doi-asserted-by":"publisher","first-page":"532","DOI":"10.1890\/0012-9658(2000)081[0532:COAAGT]2.0.CO;2","volume":"81","author":"CG Eckert","year":"2000","unstructured":"Eckert, C. G. Contributions of autogamy and geitonogamy to self-fertilization in a mass-flowering, clonal plant. Ecology 81, 532\u2013542 (2000).","journal-title":"Ecology"},{"key":"29571_CR59","doi-asserted-by":"publisher","first-page":"1503","DOI":"10.3732\/ajb.92.9.1503","volume":"92","author":"JW Busch","year":"2005","unstructured":"Busch, J. W. The evolution of self-compatibility in geographically peripheral populations of Leavenworthia alabamica (Brassicaceae). Am. J. Bot. 92, 1503\u20131512 (2005).","journal-title":"Am. J. Bot."},{"key":"29571_CR60","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1007\/BF00937433","volume":"184","author":"AO Olivencia","year":"1993","unstructured":"Olivencia, A. O. & Alcaraz, J. A. D. Sexual reproduction in somescrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands. Plant. Syst. Evol. 184, 159\u2013174 (1993).","journal-title":"Plant. Syst. Evol."},{"key":"29571_CR61","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1080\/23818107.2019.1652847","volume":"166","author":"ML Navarro-P\u00e9rez","year":"2019","unstructured":"Navarro-P\u00e9rez, M. L., L\u00f3pez, J., Rodr\u00edguez-Ria\u00f1o, T. & Ortega-Olivencia, A. Reproductive system of two mediterranean Scrophularia species with large, showy flowers. Bot. Lett. 166, 467\u2013477 (2019).","journal-title":"Bot. Lett."},{"key":"29571_CR62","doi-asserted-by":"publisher","DOI":"10.3791\/54663","author":"BA Pace","year":"2016","unstructured":"Pace, B. A., Alexander, H. M., Emry, D. J. & Mercer, K. L. Reliable method for assessing seed germination, dormancy, and mortality under field conditions. J. Vis. Exp. https:\/\/doi.org\/10.3791\/54663 (2016).","journal-title":"J. Vis. Exp."},{"key":"29571_CR63","doi-asserted-by":"publisher","first-page":"30","DOI":"10.1038\/hdy.2010.151","volume":"107","author":"JS Miller","year":"2011","unstructured":"Miller, J. S. & Kostyun, J. L. Functional gametophytic self-incompatibility in a peripheral population of Solanum peruvianum (Solanaceae). Heredity 107, 30\u201339 (2011).","journal-title":"Heredity"},{"key":"29571_CR64","doi-asserted-by":"publisher","first-page":"1279","DOI":"10.1111\/eva.12933","volume":"13","author":"E Durand","year":"2020","unstructured":"Durand, E. et al. Evolution of self-incompatibility in the brassicaceae: lessons from a textbook example of natural selection. Evol. Appl. 13, 1279\u20131297 (2020).","journal-title":"Evol. Appl."},{"key":"29571_CR65","doi-asserted-by":"publisher","first-page":"e12910","DOI":"10.1111\/csp2.12910","volume":"5","author":"CAT Doyle","year":"2023","unstructured":"Doyle, C. A. T. et al. Reproductive characteristics, population genetics, and pairwise kinship inform strategic recovery of a plant species in a fragmented landscape. Conserv. Sci. Pract. 5, e12910 (2023).","journal-title":"Conserv. Sci. Pract."},{"key":"29571_CR66","doi-asserted-by":"publisher","first-page":"100719","DOI":"10.1016\/j.xplc.2023.100719","volume":"5","author":"D Zhang","year":"2024","unstructured":"Zhang, D. et al. Molecular insights into self-incompatibility systems: from evolution to breeding. Plant. Commun. 5, 100719 (2024).","journal-title":"Plant. Commun."},{"key":"29571_CR67","doi-asserted-by":"publisher","first-page":"380","DOI":"10.1038\/347380a0","volume":"347","author":"B Charlesworth","year":"1990","unstructured":"Charlesworth, B., Charlesworth, D. & Morgan, M. T. Genetic loads and estimates of mutation rates in highly inbred plant populations. Nature 347, 380\u2013382 (1990).","journal-title":"Nature"},{"key":"29571_CR68","doi-asserted-by":"publisher","first-page":"370","DOI":"10.1086\/297041","volume":"153","author":"DG Lloyd","year":"1992","unstructured":"Lloyd, D. G. Self- and Cross-Fertilization in Plants. II. The selection of Self- fertilization. Int. J. Plant. Sci. 153, 370\u2013380 (1992).","journal-title":"Int. J. Plant. Sci."},{"key":"29571_CR69","doi-asserted-by":"publisher","first-page":"407","DOI":"10.3389\/fpls.2019.00407","volume":"10","author":"H Claessen","year":"2019","unstructured":"Claessen, H., Keulemans, W., Van De Poel, B. & De Storme, N. Finding a compatible partner: Self-Incompatibility in European Pear (Pyrus communis); molecular Control, genetic Determination, and impact on fertilization and fruit set. Front. Plant. Sci. 10, 407 (2019).","journal-title":"Front. Plant. Sci."},{"key":"29571_CR70","doi-asserted-by":"publisher","DOI":"10.1093\/aob\/mcaf047","author":"S Chabert","year":"2025","unstructured":"Chabert, S. & Mallinger, R. E. Self-sterility, self-incompatibility and xenia: a review of the mechanisms of cross-pollination benefits in animal-pollinated crops. Ann. Bot. https:\/\/doi.org\/10.1093\/aob\/mcaf047 (2025).","journal-title":"Ann. Bot."},{"key":"29571_CR71","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1093\/aob\/mcq149","volume":"106","author":"G Vaughton","year":"2010","unstructured":"Vaughton, G., Ramsey, M. & Johnson, S. D. Pollination and late-acting self-incompatibility in Cyrtanthus breviflorus (Amaryllidaceae): implications for seed production. Ann. Bot. 106, 547\u2013555 (2010).","journal-title":"Ann. Bot."},{"key":"29571_CR72","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1007\/s004970100068","volume":"13","author":"R Aguilar","year":"2001","unstructured":"Aguilar, R. & Bernardello, G. The breeding system of lycium cestroides: a Solanaceae with ovarian self-incompatibility. Sex. Plant. Reprod. 13, 273\u2013277 (2001).","journal-title":"Sex. Plant. Reprod."},{"key":"29571_CR73","doi-asserted-by":"publisher","first-page":"926","DOI":"10.3390\/plants12040926","volume":"12","author":"M Ferriol","year":"2023","unstructured":"Ferriol, M. et al. Pre- and Post-Zygotic barriers contribute to reproductive isolation and correlate with genetic distance in cucumis. Plants 12, 926 (2023).","journal-title":"Plants"},{"key":"29571_CR74","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1086\/700314","volume":"179","author":"\u00d8H Opedal","year":"2018","unstructured":"Opedal, \u00d8. H. Herkogamy, a principal functional trait of plant reproductive biology. Int. J. Plant. Sci. 179, 677\u2013687 (2018).","journal-title":"Int. J. Plant. Sci."},{"key":"29571_CR75","doi-asserted-by":"publisher","first-page":"529","DOI":"10.1098\/rstb.2009.0226","volume":"365","author":"LD Harder","year":"2010","unstructured":"Harder, L. D. & Aizen, M. A. Floral adaptation and diversification under pollen limitation. Philos. Trans. R Soc. B Biol. Sci. 365, 529\u2013543 (2010).","journal-title":"Philos. Trans. R Soc. B Biol. Sci."},{"key":"29571_CR76","doi-asserted-by":"publisher","first-page":"plw061","DOI":"10.1093\/aobpla\/plw061","volume":"8","author":"\u00c5 Lankinen","year":"2016","unstructured":"Lankinen, \u00c5. & Strandh, M. Differential selection on pollen and pistil traits in relation to pollencompetition in the context of a sexual conflict over timing of stigmareceptivity. AoB Plants. 8, plw061 (2016).","journal-title":"AoB Plants"},{"key":"29571_CR77","doi-asserted-by":"publisher","first-page":"1237","DOI":"10.3732\/ajb.89.8.1237","volume":"89","author":"HJ Young","year":"2002","unstructured":"Young, H. J. & Gravitz, L. The effects of stigma age on receptivity in Silene alba (Caryophyllaceae). Am. J. Bot. 89, 1237\u20131241 (2002).","journal-title":"Am. J. Bot."},{"key":"29571_CR78","doi-asserted-by":"crossref","unstructured":"Brito, M. S. et al. Pollination triggers female gametophyte development in immature Nicotiana tabacum flowers. Front Plant. Sci 6\u00a0(2015).","DOI":"10.3389\/fpls.2015.00561"},{"key":"29571_CR79","doi-asserted-by":"publisher","first-page":"1205816","DOI":"10.3389\/fpls.2023.1205816","volume":"14","author":"LT Tran","year":"2023","unstructured":"Tran, L. T., Sugimoto, K., Kasozi, M., Mitalo, O. W. & Ezura, H. Pollination, pollen tube growth, and fertilization independently contribute to fruit set and development in tomato. Front. Plant. Sci. 14, 1205816 (2023).","journal-title":"Front. Plant. Sci."},{"key":"29571_CR80","doi-asserted-by":"publisher","first-page":"865","DOI":"10.3390\/biology14070865","volume":"14","author":"U Layek","year":"2025","unstructured":"Layek, U., Majhi, P., Das, A., Karmakar, P. & Kundu, A. Seasonal variation in flower traits, visitor traits, and reproductive success of Solanum sisymbriifolium Lamarck (Solanaceae) in the Rarh region of West Bengal, India. Biology 14, 865 (2025).","journal-title":"Biology"},{"key":"29571_CR81","doi-asserted-by":"publisher","first-page":"115","DOI":"10.21273\/HORTSCI.30.1.115","volume":"30","author":"AA Abdul-Baki","year":"1995","unstructured":"Abdul-Baki, A. A. & Stommel, J. R. Pollen viability and fruit set of tomato genotypes under optimumand High-temperature regimes. HortScience 30, 115\u2013117 (1995).","journal-title":"HortScience"},{"key":"29571_CR82","first-page":"26","volume":"38","author":"C Manjunatha","year":"2010","unstructured":"Manjunatha, C., Tomar, B. S., Balraj, S. & Kalyanrao, P. Effects of method and time of pollination on tomato (Lycopersicon esculantum) fruit set and seed quality. Seed Res. 38, 26\u201330 (2010).","journal-title":"Seed Res."},{"key":"29571_CR83","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1046\/j.1365-3040.2000.00589.x","volume":"23","author":"S Sato","year":"2000","unstructured":"Sato, S., Peet, M. M. & Thomas, J. F. Physiological factors limit fruit set of tomato (Lycopersicon esculentum Mill.) under chronic, mild heat stress. Plant. Cell. Environ. 23, 719\u2013726 (2000).","journal-title":"Plant. Cell. Environ."},{"key":"29571_CR84","doi-asserted-by":"publisher","first-page":"1187","DOI":"10.1093\/jexbot\/53.371.1187","volume":"53","author":"S Sato","year":"2002","unstructured":"Sato, S., Peet, M. M. & Thomas, J. F. Determining critical pre- and post\u2010anthesis periods and physiological processes in Lycopersicon esculentum Mill. exposed to moderately elevated temperatures. J. Exp. Bot. 53, 1187\u20131195 (2002).","journal-title":"J. Exp. Bot."},{"key":"29571_CR85","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1046\/j.0016-8025.2001.00807.x","volume":"25","author":"AN Erickson","year":"2002","unstructured":"Erickson, A. N. & Markhart, A. H. Flower developmental stage and organ sensitivity of bell pepper (Capsicum annuum L.) to elevated temperature. Plant. Cell. Environ. 25, 123\u2013130 (2002).","journal-title":"Plant. Cell. Environ."},{"key":"29571_CR86","doi-asserted-by":"publisher","first-page":"1495610","DOI":"10.3389\/fsufs.2024.1495610","volume":"8","author":"M Erfatpour","year":"2024","unstructured":"Erfatpour, M., MacLean, D., Lahlali, R. & Jiang, Y. Ovule and seed development of crop plants in response to climate change. Front. Sustain. Food Syst. 8, 1495610 (2024).","journal-title":"Front. Sustain. Food Syst."},{"key":"29571_CR87","doi-asserted-by":"publisher","first-page":"968","DOI":"10.1111\/j.1461-0248.2006.00927.x","volume":"9","author":"R Aguilar","year":"2006","unstructured":"Aguilar, R., Ashworth, L., Galetto, L. & Aizen, M. A. Plant reproductive susceptibility to habitat fragmentation: review and synthesis through a meta-analysis. Ecol. Lett. 9, 968\u2013980 (2006).","journal-title":"Ecol. Lett."},{"key":"29571_CR88","doi-asserted-by":"publisher","first-page":"599","DOI":"10.1016\/j.tplants.2009.08.005","volume":"14","author":"AT Kramer","year":"2009","unstructured":"Kramer, A. T. & Havens, K. Plant conservation genetics in a changing world. Trends Plant. Sci. 14, 599\u2013607 (2009).","journal-title":"Trends Plant. Sci."},{"key":"29571_CR89","doi-asserted-by":"publisher","first-page":"2706","DOI":"10.1002\/ece3.2897","volume":"7","author":"L Sol\u00eds-Montero","year":"2017","unstructured":"Sol\u00eds-Montero, L. & Vallejo\u2010Mar\u00edn, M. Does the morphological fit between flowers and pollinators affect pollen deposition? An experimental test in a buzz\u2010pollinated species with anther dimorphism. Ecol. Evol. 7, 2706\u20132715 (2017).","journal-title":"Ecol. Evol."},{"key":"29571_CR90","doi-asserted-by":"publisher","first-page":"1603","DOI":"10.3390\/agronomy11081603","volume":"11","author":"ML Alcaraz","year":"2021","unstructured":"Alcaraz, M. L. & Hormaza, J. I. Fruit set in avocado: pollen Limitation, pollen load Size, and selective fruit abortion. Agronomy 11, 1603 (2021).","journal-title":"Agronomy"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29571-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29571-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29571-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,5]],"date-time":"2026-01-05T14:27:10Z","timestamp":1767623230000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29571-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,5]]},"references-count":90,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["29571"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-29571-5","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,5]]},"assertion":[{"value":"27 August 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 November 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"5 January 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"302"}}