{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T08:52:17Z","timestamp":1791276737680,"version":"4.1.0"},"reference-count":60,"publisher":"MDPI AG","issue":"19","license":[{"start":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T00:00:00Z","timestamp":1791244800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100006769","name":"Russian Science Foundation","doi-asserted-by":"publisher","award":["26-16-00147"],"award-info":[{"award-number":["26-16-00147"]}],"id":[{"id":"10.13039\/501100006769","id-type":"DOI","asserted-by":"publisher"}]},{"award":["26-16-00147"],"award-info":[{"award-number":["26-16-00147"]}],"id":[{"id":"https:\/\/ror.org\/03y2gwe85","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Distant hybridization is an important breeding strategy that can effectively expand the genetic pool for developing new, high-yielding, and stress-resistant cultivars. However, reproductive isolation often hinders the application of this approach. Intergeneric reproductive barriers (IRBs) manifest during pollen\u2013pistil interactions in plants belonging to different genera, yet the mechanisms underlying these barriers remain largely unknown. In this study, we investigated pollen\u2013pistil interactions during the progamic phase of fertilization in three Solanaceae species using several intergeneric cross combinations. Pollen of all tested species germinated on the stigma surface, grew through the stylar transmitting tissue, and ceased elongation at a distance of 300\u2013920 \u00b5m therefrom due to the action of a reproductive barrier. We suggest that programmed cell death (PCD) is a factor contributing to IRBs in Solanaceae. The obtained data indicate that, in all cross combinations, the rejection of incompatible pollen tubes (PTs) is accompanied by increased activity of caspase-like proteases (CLPs), as confirmed by in vivo visualization of active CLPs and spectrophotometric measurement of their activity. Specifically, incompatible PT growth in stylar tissues during the first 4 h after pollination was accompanied by a five- to six-fold increase in caspase-3-like protease activity. Notably, the arrest of PT growth in distant hybridization shows striking similarity to that observed during S-RNase-based self-incompatibility (SI) in Solanaceae. We anticipate that further research will ultimately provide insights into the regulation of reproductive barriers during distant hybridization.<\/jats:p>","DOI":"10.3390\/plants15193048","type":"journal-article","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T08:06:47Z","timestamp":1791274007000},"page":"3048","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Caspase-Like Proteases as Contributors in the Functioning of Reproductive Barriers During Distant Hybridization in Solanaceae"],"prefix":"10.3390","volume":"15","author":[{"given":"Alexej I.","family":"Ulianov","sequence":"first","affiliation":[{"name":"All-Russia Research Institute of Agricultural Biotechnology, 127550 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1299-798X","authenticated-orcid":false,"given":"Yaroslav Yu.","family":"Golivanov","sequence":"additional","affiliation":[{"name":"All-Russia Research Institute of Agricultural Biotechnology, 127550 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-2679-424X","authenticated-orcid":false,"given":"Tatiana P.","family":"Molchanova","sequence":"additional","affiliation":[{"name":"All-Russia Research Institute of Agricultural Biotechnology, 127550 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6600-1693","authenticated-orcid":false,"given":"Oksana A.","family":"Muratova","sequence":"additional","affiliation":[{"name":"All-Russia Research Institute of Agricultural Biotechnology, 127550 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Natalia A.","family":"Milyukova","sequence":"additional","affiliation":[{"name":"All-Russia Research Institute of Agricultural Biotechnology, 127550 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ekaterina V.","family":"Zakharova","sequence":"additional","affiliation":[{"name":"All-Russia Research Institute of Agricultural Biotechnology, 127550 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,10,6]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"148","DOI":"10.20546\/ijcmas.2022.1105.019","article-title":"Concept of gene pool and distant hybridization in vegetable crops","volume":"11","author":"Subhasmita","year":"2022","journal-title":"Int. J. Curr. Microbiol. Appl. Sci."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Song, H., Zou, T., Gong, S., Zhang, C., Sun, X., Luo, Y., and Sun, X. (2025). Research on the creation and application of a doubled haploid for distant hybrid pumpkin. Hortic. Plant J., in press.","DOI":"10.1016\/j.hpj.2024.10.001"},{"key":"ref_3","unstructured":"Roy, S.C., and Shil, P. (2020). Black rice developed through interspecific hybridization (O. sativa \u00d7 O. rufipogon): Origin of black rice gene from Indian wild rice. bioRxiv."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"932737","DOI":"10.3389\/fpls.2022.932737","article-title":"Interspecific hybridization is an important driving force for origin and diversification of Asian cultivated rice Oryza sativa L.","volume":"13","author":"Zhou","year":"2022","journal-title":"Front. Plant Sci."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Tan, M., Chen, R., Chen, X., Shahid, M.Q., Liu, X., and Wu, J. (2023). In vitro induction of interspecific hybrid and polyploidy derived from Oryza officinalis Wall. Plants, 12.","DOI":"10.3390\/plants12163001"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"6321","DOI":"10.1007\/s10722-025-02402-2","article-title":"Potential of intersubspecific hybrids in rice: An overview of sterility loci and overcoming strategies","volume":"72","author":"Saravanan","year":"2025","journal-title":"Genet. Resour. Crop Evol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1186\/s13059-019-1744-x","article-title":"Frequent intra- and inter-species introgression shapes the landscape of genetic variation in bread wheat","volume":"20","author":"Cheng","year":"2019","journal-title":"Genome Biol."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Han, G.H., Wang, J., Yan, H.W., Cao, L.J., Liu, S.Y., Li, X.Q., Zhou, Y.L., Liu, W., Gu, T.T., and Shi, Z.P. (2023). Development and molecular cytogenetic identification of a new wheat-rye 6RL ditelosomic addition and 1R (1B) substitution line with powdery mildew resistance. J. Integr. Agric., in press.","DOI":"10.1016\/j.jia.2023.10.004"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Pang, J., Huang, C., Wang, Y., Wen, X., Deng, P., Li, T., Wang, C., Liu, X., Chen, C., and Zhao, J. (2023). Molecular cytological analysis and specific marker development in wheat-Psathyrostachys huashanica Keng 3Ns additional line with elongated glume. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24076726"},{"key":"ref_10","first-page":"e190007","article-title":"Interspecific hybridization for Brassica crop improvement","volume":"1","author":"Katche","year":"2019","journal-title":"Crop Breed. Genet. Genom."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"109412","DOI":"10.1016\/j.scienta.2020.109412","article-title":"Production of allohexaploid Brassica hybrid between tuber mustard (Brassica juncea L. var. crassicaulis Chen & Yang) and Chinese kale (Brassica oleracea var. alboglabra Bailey)","volume":"270","author":"Chen","year":"2020","journal-title":"Sci. Hortic."},{"key":"ref_12","first-page":"2022","article-title":"Interspecific hybridization and plant breeding: From historical retrospective through work of Mendel to current crops","volume":"11","year":"2022","journal-title":"Plants"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"110801","DOI":"10.1016\/j.scienta.2021.110801","article-title":"Exotic genes for solving emerging peach production challenges","volume":"295","author":"Gradziel","year":"2022","journal-title":"Sci. Hortic."},{"key":"ref_14","first-page":"22","article-title":"A review on application of mechanism promoting self and cross pollination in crop plants","volume":"16","author":"Mudhalvan","year":"2024","journal-title":"Biol. Forum\u2014Int. J."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"112346","DOI":"10.1016\/j.scienta.2023.112346","article-title":"Effects of species and varietal differences on intergeneric cross-compatibility between apple and pear","volume":"321","author":"Morimoto","year":"2023","journal-title":"Sci. Hortic."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1007\/s00425-023-04265-2","article-title":"Characterization of pollen tube development in distant hybridization of Chinese cork oak (Quercus variabilis L.)","volume":"258","author":"Ke","year":"2023","journal-title":"Planta"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1038\/s41420-024-01821-x","article-title":"Ac-DEVD-CHO (caspase-3\/DEVDase inhibitor) suppresses self-incompatibility\u2013induced programmed cell death in the pollen tubes of petunia (Petunia hybrida E. Vilm.)","volume":"10","author":"Zakharova","year":"2024","journal-title":"Cell Death Discov."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Zakharova, E., Khanina, T., Knyazev, A., Milyukova, N., and Kovaleva, L.V. (2023). Hormonal signaling during dPCD: Cytokinin as the determinant of RNase-based self-incompatibility in Solanaceae. Biomolecules, 13.","DOI":"10.3390\/biom13071033"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1007\/s00709-019-01430-x","article-title":"Aminooxyacetic acid (AOA), inhibitor of 1-aminocyclopropane-1-carboxylic acid (ACC) synthesis, suppresses self-incompatibility-induced programmed cell death in self-incompatible Petunia hybrida L. pollen tubes","volume":"257","author":"Kovaleva","year":"2020","journal-title":"Protoplasma"},{"key":"ref_20","first-page":"40","article-title":"Ac-DEVD-CHO (caspase-3\/DEVDase inhibitor) suppresses self-incompatibility-induced programmed cell death in the pollen tubes of Solanum pennellii Correll","volume":"8","author":"Molchanova","year":"2025","journal-title":"Ecobiotech"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1007\/s00425-020-03526-8","article-title":"Glycosylphosphatidylinositol-anchored proteins mediate the interactions between pollen\/pollen tube and pistil tissues","volume":"253","author":"Yu","year":"2021","journal-title":"Planta"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"687","DOI":"10.1111\/nph.15645","article-title":"Pollen tube integrity regulation in flowering plants: Insights from molecular assemblies on the pollen tube surface","volume":"222","author":"Ge","year":"2019","journal-title":"New Phytol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Bordeleau, S.J., Canales Sanchez, L.E., and Goring, D.R. (2022). Finding new Arabidopsis receptor kinases that regulate compatible pollen-pistil interactions. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.1022684"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"558","DOI":"10.1104\/pp.19.01505","article-title":"Durotropic growth of pollen tubes","volume":"183","author":"Reimann","year":"2020","journal-title":"Plant Physiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1038\/s41586-022-05640-x","article-title":"Stigma receptors control intraspecies and interspecies barriers in Brassicaceae","volume":"614","author":"Huang","year":"2023","journal-title":"Nature"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Zheng, S., Wang, F., Liu, Z., Zhang, H., Zhang, L., and Chen, D. (2024). The role of female and male genes in regulating pollen tube guidance in flowering plants. Genes, 15.","DOI":"10.3390\/genes15111367"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Callaway, T.D., and Singh-Cundy, A. (2019). HD-AGPs as speciation genes: Positive selection on a proline-rich domain in non-hybridizing species of Petunia, Solanum, and Nicotiana. Plants, 8.","DOI":"10.3390\/plants8070211"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1093\/genetics\/22.1.130","article-title":"Cytological phenomena connected with self-sterility in the flowering plants","volume":"22","author":"Sears","year":"1937","journal-title":"Genetics"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1104\/pp.16.01286","article-title":"Pollen-pistil interactions and their role in mate selection","volume":"173","author":"Bedinger","year":"2017","journal-title":"Plant Physiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1093\/plphys\/kiae050","article-title":"Molecular basis and evolutionary drivers of endosperm-based hybridization barriers","volume":"195","author":"Bente","year":"2024","journal-title":"Plant Physiol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"koag031","DOI":"10.1093\/plcell\/koag031","article-title":"Papaver S-determinants trigger mitochondrially derived ROS production and disrupt energy metabolism in incompatible pollen tubes","volume":"38","author":"Wang","year":"2026","journal-title":"Plant Cell"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"4692","DOI":"10.1093\/plcell\/koae245","article-title":"Regulating programmed cell death in plant cells: Intracellular acidification plays a pivotal role together with calcium signaling","volume":"36","author":"Bosch","year":"2024","journal-title":"Plant Cell"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1038\/367563a0","article-title":"S-RNase expressed in transgenic Nicotiana causes S-allele-specific pollen rejection","volume":"367","author":"Murfett","year":"1994","journal-title":"Nature"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"14005","DOI":"10.1038\/nplants.2014.5","article-title":"Gene duplication and genetic exchange drive the evolution of S-RNase-based self-incompatibility in Petunia","volume":"1","author":"Kubo","year":"2015","journal-title":"Nat. Plants"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1094","DOI":"10.1111\/j.1365-313X.2008.03487.x","article-title":"Identification of major lysine residues of S3-RNase of Petunia inflata involved in ubiquitin-26S proteasome-mediated degradation in vitro","volume":"54","author":"Hua","year":"2008","journal-title":"Plant J."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1146\/annurev-arplant-042809-112256","article-title":"The cullin-RING ubiquitin-protein ligases","volume":"62","author":"Hua","year":"2011","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1016\/S1055-7903(03)00195-7","article-title":"S-RNases and sexual incompatibility: Structure, functions, and evolutionary perspectives","volume":"29","author":"Roalson","year":"2003","journal-title":"Mol. Phylogenet. Evol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1069","DOI":"10.1093\/jxb\/erp024","article-title":"Darwin\u2019s foundation for investigating self-incompatibility and the progress toward a physiological model for S-RNase-based SI","volume":"60","author":"McClure","year":"2009","journal-title":"J. Exp. Bot."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.plaphy.2020.11.052","article-title":"The regulatory framework of developmentally programmed cell death in floral organs: A review","volume":"158","author":"Wang","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"4749","DOI":"10.1093\/jxb\/err196","article-title":"Classes of programmed cell death in plants, compared to those in animals","volume":"62","year":"2011","journal-title":"J. Exp. Bot."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1074\/jbc.M304468200","article-title":"Ultraviolet-C overexposure induces programmed cell death in Arabidopsis, which is mediated by caspase-like activities and which can be suppressed by caspase inhibitors, p35 and Defender against Apoptotic Death","volume":"279","author":"Danon","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1038\/sj.cdd.4401330","article-title":"VEIDase is a principal caspase-like activity involved in plant programmed cell death and essential for embryonic pattern formation","volume":"11","author":"Bozhkov","year":"2004","journal-title":"Cell Death Differ."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Zhang, Q.F., Li, J., Bi, F.C., Liu, Z., Chang, Z.Y., Wang, L.Y., Huang, L.Q., and Yao, N. (2020). Ceramide-induced cell death depends on calcium and caspase-like activity in rice. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.00145"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1093\/jxb\/erm352","article-title":"What happened to plant caspases?","volume":"59","author":"Bonneau","year":"2008","journal-title":"J. Exp. Bot."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1038\/s41477-017-0020-x","article-title":"Reassessing apoptosis in plants","volume":"3","author":"Dickman","year":"2017","journal-title":"Nat. Plants"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1129","DOI":"10.1016\/S0960-9822(98)70469-5","article-title":"Caspases and programmed cell death in the hypersensitive response of plants to pathogens","volume":"8","author":"Lam","year":"1998","journal-title":"Curr. Biol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1007\/s00709-020-01587-w","article-title":"Caspase-like proteases and the phytohormone cytokinin as determinants of S-RNase\u2013based self-incompatibility\u2013induced PCD in Petunia hybrida L.","volume":"258","author":"Zakharova","year":"2021","journal-title":"Protoplasma"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1083\/jcb.200604011","article-title":"Actin depolymerization is sufficient to induce programmed cell death in self-incompatible pollen","volume":"174","author":"Thomas","year":"2006","journal-title":"J. Cell Biol."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Liu, W., Fan, J., Li, J., Song, Y., Li, Q., Zhang, Y.E., and Xue, Y. (2014). SCFSLF-mediated cytosolic degradation of S-RNase is required for cross-pollen compatibility in S-RNase-based self-incompatibility in Petunia hybrida. Front. Genet., 5.","DOI":"10.3389\/fgene.2014.00228"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Zakharova, E.V., Ulianov, A.I., Golivanov, Y.Y., Molchanova, T.P., Orlova, Y.V., and Muratova, O.A. (2025). Pollen\u2013pistil interaction during distant hybridization in plants. Agronomy, 15.","DOI":"10.3390\/agronomy15071732"},{"key":"ref_51","first-page":"1234","article-title":"Dominance modifiers at the Arabidopsis self-incompatibility locus retain proto-miRNA features and act through non-canonical pathways","volume":"15","author":"Batista","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1093\/jxb\/erg045","article-title":"S-RNase complexes and pollen rejection","volume":"54","author":"Hancock","year":"2003","journal-title":"J. Exp. Bot."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1964","DOI":"10.3732\/ajb.1600356","article-title":"Interspecific reproductive barriers between sympatric populations of wild tomato species (Solanum section Lycopersicon)","volume":"103","author":"Baek","year":"2016","journal-title":"Am. J. Bot."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"718","DOI":"10.1111\/tpj.13416","article-title":"HT proteins contribute to S-RNase-independent pollen rejection in Solanum","volume":"89","author":"Lu","year":"2017","journal-title":"Plant J."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"927","DOI":"10.1093\/plphys\/kiad301","article-title":"Stop and go signals at the stigma\u2013pollen interface of the Brassicaceae","volume":"193","author":"Nasrallah","year":"2023","journal-title":"Plant Physiol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1111\/nph.15340","article-title":"Correlated evolution of self and interspecific incompatibility across the range of a Texas wildflower","volume":"221","author":"Roda","year":"2019","journal-title":"New Phytol."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"de Nettancourt, D. (2001). The genetics of self-incompatibility. Incompatibility and Incongruity in Wild and Cultivated Plants, Springer.","DOI":"10.1007\/978-3-662-04502-2"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1002","DOI":"10.3732\/ajb.1200616","article-title":"Interactions of hybridization and mating systems: A case study in Leptosiphon (Polemoniaceae)","volume":"100","author":"Goodwillie","year":"2013","journal-title":"Am. J. Bot."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Ulianov, A.I., Golivanov, Y.Y., Molchanova, T.P., Orlova, Y.V., Muratova, O.A., Milyukova, N.A., and Zakharova, E.V. (2026). Distant Hybridization in Solanaceae: Similarity to Self-Incompatibility of Pollen Tube Growth Arrest as a Reproductive Barrier. Agronomy, 16.","DOI":"10.3390\/agronomy16141326"},{"key":"ref_60","first-page":"146","article-title":"A simple colorimetric method for determination of protein","volume":"131","author":"Bramhall","year":"1983","journal-title":"Anal. Biochem."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/19\/3048\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T08:17:48Z","timestamp":1791274668000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/19\/3048"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,6]]},"references-count":60,"journal-issue":{"issue":"19","published-online":{"date-parts":[[2026,10]]}},"alternative-id":["plants15193048"],"URL":"https:\/\/doi.org\/10.3390\/plants15193048","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,6]]}}}