[1]

He N, Zhang C, Qi X, Zhao S, Tao Y, et al. 2013. Draft genome sequence of the mulberry tree Morus notabilis. Nature Communications 4:2445

doi: 10.1038/ncomms3445
[2]

Lin CY, Lay HL. 2013. Characteristics of fruit growth, component analysis and antioxidant activity of mulberry (Morus spp.) Scientia Horticulturae 162:285−292

doi: 10.1016/j.scienta.2013.08.009
[3]

Tian S, Wang M, Liu C, Zhao H, Zhao B. 2019. Mulberry leaf reduces inflammation and insulin resistance in type 2 diabetic mice by TLRs and insulin Signalling pathway. BMC Complementary and Alternative Medicine 19:326

doi: 10.1186/s12906-019-2742-y
[4]

Zhang F, Liu X, Xia H, Wu H, Zong Y, et al. 2025. Identification of genetic loci for growth and stem form traits in hybrid Liriodendron via a genome-wide association study. Forestry Research 5:e001

doi: 10.48130/forres-0025-0001
[5]

Gong B, Gao J, Xie Y, Zhang H, Zhu W, et al. 2024. Development of wheat-tetraploid Thinopyrum elongatum 4EL small fragment translocation lines with stripe rust resistance gene Yr4EL. Theoretical and Applied Genetics 137:246

doi: 10.1007/s00122-024-04756-0
[6]

Guan H, Huang B, Yan X, Zhao J, Yang S, et al. 2024. Identification of distinct roses suitable for future breeding by phenotypic and genotypic evaluations of 192 rose germplasms. Horticulture Advances 2:5

doi: 10.1007/s44281-023-00024-1
[7]

Chen X, Gao Y, Zhang D, Gao Y, Song Y, et al. 2023. Evaluation of salinity resistance and combining ability analysis in the seedlings of mulberry hybrids (Morus alba L.). Physiology and Molecular Biology of Plants 29:543−557

doi: 10.1007/s12298-023-01304-w
[8]

Britten EJ. 1960. A proposed classification of screening methods for plant breeding programs. Euphytica 9:293−303

doi: 10.1007/BF00029481
[9]

Tani N, Kawahara T, Yoshimaru H, Hoshi Y. 2003. Development of SCAR markers distinguishing pure seedlings of the endangered species Morus boninensis from M. boninensis × M. acidosa hybrids for conservation in Bonin (Ogasawara) Islands. Conservation Genetics 4:605−612

doi: 10.1023/A:1025655331429
[10]

Stetter MG, Zeitler L, Steinhaus A, Kroener K, Biljecki M, et al. 2016. Crossing methods and cultivation conditions for rapid production of segregating populations in three grain amaranth species. Frontiers in Plant Science 7:816

doi: 10.3389/fpls.2016.00816
[11]

Du F, Jiang J, Jia H, Zhao XY, Wang WH, et al. 2015. Selection of generally applicable SSR markers for evaluation of genetic diversity and identity in Lilium. Biochemical Systematics and Ecology 61:278−285

doi: 10.1016/j.bse.2015.05.002
[12]

Yin ZF, Zhao B, Bi WL, Chen L, Wang QC. 2013. Direct shoot regeneration from basal leaf segments of Lilium and assessment of genetic stability in regenerants by ISSR and AFLP markers. In Vitro Cellular & Developmental Biology - Plant 49:333−342

doi: 10.1007/s11627-013-9501-4
[13]

Xuan Y, Wu Y, Li P, Liu R, Luo Y, et al. 2019. Molecular phylogeny of mulberries reconstructed from ITS and two cpDNA sequences. PeerJ 7:e8158

doi: 10.7717/peerj.8158
[14]

Zeng Q, Chen H, Zhang C, Han M, Li T, et al. 2015. Definition of eight mulberry species in the genus Morus by internal transcribed spacer-based phylogeny. PLoS One 10:e0135411

doi: 10.1371/journal.pone.0135411
[15]

Mandák B, Krak K, Vít P, Lomonosova MN, Belyayev A, et al. 2018. Hybridization and polyploidization within the Chenopodium album aggregate analysed by means of cytological and molecular markers. Molecular Phylogenetics and Evolution 129:189−201

doi: 10.1016/j.ympev.2018.08.016
[16]

Moarefi N, Michael P, Beckett P, Nkongolo K. 2018. Identification of molecular markers differentiating Betula papyrifera and B. pumila populations from Northern Ontario (Canada). American Journal of Environmental Sciences 14:246−256

doi: 10.3844/ajessp.2018.246.256
[17]

Clevenger J, Chavarro C, Pearl SA, Ozias-Akins P, Jackson SA. 2015. Single nucleotide polymorphism identification in polyploids: a review, example, and recommendations. Molecular Plant 8:831−846

doi: 10.1016/j.molp.2015.02.002
[18]

Wang H, Liu J, Xu X, Li Y, Yuan J, et al. 2024. MGCPdb, a collective resource for mulberry genome size, chromosome number, and ploidy. Forestry Research 4:e027

doi: 10.48130/forres-0024-0024
[19]

Wang M, Zhu M, Qian J, Yang Z, Shang F, et al. 2024. Phylogenomics of mulberries (Morus, Moraceae) inferred from plastomes and single copy nuclear genes. Molecular Phylogenetics and Evolution 197:108093

doi: 10.1016/j.ympev.2024.108093
[20]

Xuan Y, Wang S, Li S, Yuan J, Zhou Q, et al. 2023. Chromosome constitution and genetic relationships of Morus spp. revealed by genomic in situ hybridization. BMC Plant Biology 23:428

doi: 10.1186/s12870-023-04448-9
[21]

Zeng Q, Chen M, Wang S, Xu X, Li T, et al. 2022. Comparative and phylogenetic analyses of the chloroplast genome reveal the taxonomy of the Morus genus. Frontiers in Plant Science 13:1047592

doi: 10.3389/fpls.2022.1047592
[22]

Zhao W, Pan Y, Zhang Z, Jia S, Miao X, et al. 2005. Phylogeny of the genus Morus (Urticales: Moraceae) inferred from ITS and trnL-F sequences. African Journal of Biotechnology 4:563−569

doi: 10.4314/ajb.v4i6.15143
[23]

Li X, Huang F, Chai J, Wang Q, Yu F, et al. 2021. Chromosome behavior during meiosis in pollen mother cells from Saccharum officinarum × Erianthus arundinaceus F1 hybrids. BMC Plant Biology 21:139

doi: 10.1186/s12870-021-02911-z
[24]

Park HW, Sevilleno SS, Ha MKTT, Cabahug-Braza RA, Yi JH, et al. 2024. The application of fluorescence in situ hybridization in the prescreening of Veronica hybrids. Plants 13:1264

doi: 10.3390/plants13091264
[25]

Wang G, Zhang W, Qin Y, Wu Q, Liang Q, et al. 2025. Verification of the introgression of Narenga porphyrocoma germplasm into Saccharum officinarum using molecular markers and GISH analysis. Agronomy 15:121

doi: 10.3390/agronomy15010121
[26]

Zhou M, Yong X, Zhu J, Xu Q, Liu X, et al. 2024. Chromosomal analysis of progenies between Lilium intersectional hybrids and wild species using ND-FISH and GISH. Frontiers in Plant Science 15:1461798

doi: 10.3389/fpls.2024.1461798
[27]

Kopecký D, Bartoš J, Lukaszewski AJ, Baird JH, Černoch V, et al. 2009. Development and mapping of DArT markers within the Festuca - Lolium complex. BMC Genomics 10:473

doi: 10.1186/1471-2164-10-473
[28]

De Storme N, Geelen D. 2013. Sexual polyploidization in plants–cytological mechanisms and molecular regulation. New Phytologist 198:670−684

doi: 10.1111/nph.12184
[29]

Younis A, Hwang YJ, Lim KB. 2014. Exploitation of induced 2n-gametes for plant breeding. Plant Cell Reports 33:215−223

doi: 10.1007/s00299-013-1534-y
[30]

Zhou Q, Cheng X, Kong B, Zhao Y, Li Z, et al. 2022. Heat shock-induced failure of meiosis I to meiosis II transition leads to 2n pollen formation in a woody plant. Plant Physiology 189:2110−2127

doi: 10.1093/plphys/kiac219
[31]

Dong J, Tu W, Wang H, Zuo Y, Liu T, et al. 2023. Genome sequence analysis provides insights into the mode of 2n egg formation in Solanum malmeanum. Theoretical and Applied Genetics 136:157

doi: 10.1007/s00122-023-04406-x
[32]

Palumbo F, Gabelli G, Pasquali E, Vannozzi A, Farinati S, et al. 2024. RNA-seq analyses on gametogenic tissues of alfalfa (Medicago sativa) revealed plant reproduction- and ploidy-related genes. BMC Plant Biology 24:826

doi: 10.1186/s12870-024-05542-2
[33]

Xie KD, Xia QM, Peng J, Wu XM, Xie ZZ, et al. 2019. Mechanism underlying 2n male and female gamete formation in lemon via cytological and molecular marker analysis. Plant Biotechnology Reports 13:141−149

doi: 10.1007/s11816-019-00525-4
[34]

Alix K, Gérard PR, Schwarzacher T, Heslop-Harrison JS. 2017. Polyploidy and interspecific hybridization: partners for adaptation, speciation and evolution in plants. Annals of Botany 120:183−194

doi: 10.1093/aob/mcx079
[35]

Zhong Y, Du MJ, Ji RZ, Cheng FY. 2024. Cytological origination of the first found allotriploid tree peony, Paeonia × lemoinei 'Oukan' (AAB), reveled by molecular karyotype comparation. Scientia Horticulturae 324:112563

doi: 10.1016/j.scienta.2023.112563
[36]

Parrish SB, Deng Z. 2022. Discovery and characterization of novel fertile triploids and a new chromosome number in caladium (Caladium × hortulanum). HortScience 57:1078−1085

doi: 10.21273/hortsci16701-22
[37]

Pomiès V, Turnbull N, Le Squin S, Syahputra I, Suryana E, et al. 2023. Occurrence of triploids in oil palm and their origin. Annals of Botany 131:17−32

doi: 10.1093/aob/mcac036
[38]

Kang X, Wei H. 2022. Breeding polyploid Populus: progress and perspective. Forestry Research 2:4

doi: 10.48130/FR-2022-0004
[39]

Deng H, Zhang X, Sheng S, Huang Y, Ye Z, et al. 2024. High temperature treatment induced production of unreduced 2n pollen in Camellia oleifera. Horticultural Plant Journal 10:879−896

doi: 10.1016/j.hpj.2023.10.003
[40]

Xu Y, Hu Q, Lin Q, Chi X, Shen P, et al. 2025. Polyploid crape myrtle was produced through hybridizing with 2n gametes induced by high-temperature. Euphytica 221:30

doi: 10.1007/s10681-025-03477-9
[41]

Jiang H, Jin X, Shi X, Xue Y, Jiang J, et al. 2020. Transcriptomic analysis reveals candidate genes responsive to Sclerotinia scleroterum and cloning of the Ss-inducible chitinase genes in Morus laevigata. International Journal of Molecular Sciences 21:8358

doi: 10.3390/ijms21218358
[42]

Li D, Chen G, Ma B, Zhong C, He N. 2020. Metabolic profiling and transcriptome analysis of mulberry leaves provide insights into flavonoid biosynthesis. Journal of Agricultural and Food Chemistry 68:1494−1504

doi: 10.1021/acs.jafc.9b06931
[43]

Sugiyama M, Katsube T, Koyama A, Itamura H. 2013. Varietal differences in the flavonol content of mulberry (Morus spp.) leaves and genetic analysis of quercetin 3-(6-malonylglucoside) for component breeding. Journal of Agricultural and Food Chemistry 61:9140−9147

doi: 10.1021/jf403136w
[44]

Zhou Q, Xuan Y, Li J, Qiu J, Yuan J, et al. 2025. Cytological identification of a genus-wide chromosomal fusion-fission cycles in genus Morus. Genetic Resources and Crop Evolution 72:6435−6445

doi: 10.1007/s10722-024-02312-9
[45]

Cremona G, Iovene M, Festa G, Conicella C, Parisi M. 2018. Production of embryo rescued hybrids between the landrace "Friariello" (Capsicum annuum var. annuum) and C. baccatum var. pendulum: phenotypic and cytological characterization. Euphytica 214:129

doi: 10.1007/s10681-018-2211-6
[46]

Ramzan F, Hwang YJ, Choi YH, Ramzan Y, Islam MM, et al. 2022. Evaluation of young F1 Lilium hybrids using fluorescence in situ hybridization analysis and morphological attributes. Horticulture, Environment, and Biotechnology 63:701−708

doi: 10.1007/s13580-022-00424-8
[47]

Xiao K, Sun Y, Zhou S. 2023. Revealing the abnormal meiosis and the variation of the functional female gametes of aneuploid lily (Lilium) using genomic in situ hybridization (GISH). Euphytica 219:108

doi: 10.1007/s10681-023-03238-6
[48]

Tan GX, Xiong ZY, Jin HJ, Li G, Zhu LL, et al. 2006. Characterization of interspecific hybrids between Oryza sativa L. and three wild rice species of China by genomic in situ hybridization. Journal of Integrative Plant Biology 48:1077−1083

doi: 10.1111/j.1744-7909.2006.00336.x
[49]

Zhao Y, Deng H, Chen Y, Li J, Chen S, et al. 2022. Establishment and optimization of molecular cytogenetic techniques (45S rDNA-FISH, GISH, and Fiber-FISH) in kiwifruit (Actinidia Lindl.). Frontiers in Plant Science 13:906168

doi: 10.3389/fpls.2022.906168
[50]

Sari H, Eker T, Sari D, Aksoy M, Bakır M, et al. 2023. The fastest and most reliable identification of true hybrids in the genus Pisum L. Life 13:2222

doi: 10.3390/life13112222
[51]

Meng X, Liu Z, Dai L, Zhao W, Wang J, et al. 2024. Shikimic acid accelerates phase change and flowering in Chinese jujube. Horticultural Plant Journal 10:413−424

doi: 10.1016/j.hpj.2023.02.004
[52]

Pinto RT, Abeyratne CR, Paiva LV, Benedito VA. 2023. Making headway toward enduring changes: perspectives on breeding tree crops through genome editing. Tree Genetics & Genomes 19:7

doi: 10.1007/s11295-022-01583-6
[53]

Chang H, Wang Q, Qiu Y, Qin Y, Li X, et al. 2020. Production, identification and characterization of Erianthus rockii × Narenga porphyrocoma intergeneric hybrids as a new germplasm for sugarcane breeding and genetic research. Sugar Tech 22:389−395

doi: 10.1007/s12355-020-00804-3
[54]

Xia Z, Dai X, Fan W, Liu C, Zhang M, et al. 2022. Chromosome-level genomes reveal the genetic basis of descending dysploidy and sex determination in Morus plants. Genomics Proteomics Bioinformatics 20:1119−1137

doi: 10.1016/j.gpb.2022.08.005
[55]

Venkatesh KH, Nijagunaiah R, Munirajappa. 2013. Cytogenetical studies in some diploid mulberry varieties (Moraceae). Cytologia 78:69−72

doi: 10.1508/cytologia.78.69
[56]

Venkatesh KH, Nijagunaiah R, Munirajappa. 2014. Cytogenetical studies in some triploid mulberry varieties (Moraceae). Cytologia 79:365−369

doi: 10.1508/cytologia.79.365
[57]

Dewitte A, Eeckhaut T, Van Huylenbroeck J, Van Bockstaele E. 2010. Meiotic aberrations during 2n pollen formation in Begonia. Heredity 104:215−223

doi: 10.1038/hdy.2009.111
[58]

Wang J, Li D, Shang F, Kang X. 2017. High temperature-induced production of unreduced pollen and its cytological effects in Populus. Scientific Reports 7:5281

doi: 10.1038/s41598-017-05661-x
[59]

Pang W, Wang Q, Li C, He W, Wang J, et al. 2025. ClPS1 gene-mediated manipulation of 2n pollen formation enables the creation of triploid seedless watermelon. Molecular Horticulture 5:48

doi: 10.1186/s43897-025-00170-2
[60]

Yao H, Gray AD, Auger DL, Birchler JA. 2013. Genomic dosage effects on heterosis in triploid maize. Proceedings of the National Academy of Sciences of the United States of America 110:2665−2669

doi: 10.1073/pnas.1221966110
[61]

Sun P, Nishiyama S, Asakuma H, Voorrips RE, Fu J, et al. 2021. Genomics-based discrimination of 2n gamete formation mechanisms in polyploids: a case study in nonaploid Diospyros kaki 'Akioui'. G3 Genes|Genomes|Genetics 11:jkab188

doi: 10.1093/g3journal/jkab188
[62]

Li Y, Wang Y, Wang P, Yang J, Kang X. 2016. Induction of unreduced megaspores in Eucommia ulmoides by high temperature treatment during megasporogenesis. Euphytica 212:515−524

doi: 10.1007/s10681-016-1781-4
[63]

Liu S, Sun Y, Peng M, Zhang X, Zhou S. 2024. F1 distant hybrids between two edible lilies (Lilium brownii var. viridulum and L. davidii var. unicolor) produce more n than 2n functional eggs with more recombinant chromosomes. Euphytica 220:4

doi: 10.1007/s10681-023-03269-z
[64]

Xi XJ, Jiang XB, Li D, Guo LQ, Zhang JF, et al. 2011. Induction of 2n pollen by colchicine in Populus × popularis and its triploids breeding. Silvae Genetica 60:155−160

doi: 10.1515/sg-2011-0021
[65]

Zhang X, Ren G, Li K, Zhou G, Zhou S. 2012. Genomic variation of new cultivars selected from distant hybridization in Lilium. Plant Breeding 131:227−230

doi: 10.1111/j.1439-0523.2011.01906.x
[66]

Zhao Y, Xie Y, Kong B, Cao D, Du J, et al. 2025. Genetic composition and heterozygosity of gibberellin-induced 2n gametes evaluated by SSR markers in Populus. BMC Plant Biology 25:773

doi: 10.1186/s12870-025-06824-z