{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T18:07:41Z","timestamp":1789063661543,"version":"build-2803163510"},"reference-count":44,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T00:00:00Z","timestamp":1785456000000},"content-version":"am","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"JSPS KAKENHI","award":["21J01093\/22KJ0534"],"award-info":[{"award-number":["21J01093\/22KJ0534"]}]},{"name":"JSPS KAKENHI","award":["21K15126"],"award-info":[{"award-number":["21K15126"]}]},{"name":"JSPS KAKENHI","award":["25K18486"],"award-info":[{"award-number":["25K18486"]}]},{"DOI":"10.13039\/100008608","name":"Sumitomo Foundation","doi-asserted-by":"publisher","award":["2501085"],"award-info":[{"award-number":["2501085"]}],"id":[{"id":"10.13039\/100008608","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>In angiosperms, a zygote is the initial totipotent cell that develops into an embryo. To reveal whether and how genomic and cytoplasmic contents from different cell types affect zygotic development, we analyzed developmental profiles of zygote\u2013soma hybrid cells produced by rice in vitro fertilization system. When a leaf protoplast (2\u00d7) was fused with a zygote (fusion product between egg and sperm cells; 2\u00d7), the development of most of the leaf protoplast-fused zygotes with a 1:1 gamete:soma ratio was arrested at the one-cell stage. By contrast, when a leaf protoplast (2\u00d7) was fused with a polyploid zygote (3\u00d7 or 4\u00d7), several fractions of the leaf protoplast-fused polyploid zygotes continued to develop. Furthermore, when a leaf protoplast (1\u00d7) isolated from a haploid rice plant was used for the fusion, more than half of the haploid leaf protoplast-fused zygotes developed into globular embryo-like structures and cell masses. Several of these calli then regenerated into plantlets, with the expected ploidy levels. Accordingly, leaf and zygote nuclei were integrated into the hybrid nucleus in zygote\u2013soma hybrid cells. The resulting hybrid genome was maintained during the hybrid cell development and regeneration. These findings suggest that developmental profiles of zygote\u2013soma hybrid cells are affected by the gamete:soma ratio. Furthermore, the hybrid cell fate shifts to the zygotic mode when the gamete:soma ratio changes from 1:1 to 2:1.<\/jats:p>","DOI":"10.1093\/pcp\/pcag110","type":"journal-article","created":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T11:53:35Z","timestamp":1785412415000},"source":"Crossref","is-referenced-by-count":0,"title":["Effects of somatic cell fusion on zygotic development in rice"],"prefix":"10.1093","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4786-2102","authenticated-orcid":false,"given":"Erika","family":"Toda","sequence":"first","affiliation":[{"name":"Graduate School of Agricultural and Life Sciences, The University of Tokyo , 1-1-1 Yayoi, Bunkyo, Tokyo 113-8657 ,","place":["Japan"]},{"name":"Graduate School of Science, Tokyo Metropolitan University , 1-1 Minami-osawa, Hachioji, Tokyo 192-0397 ,","place":["Japan"]},{"name":"Graduate School of Science, The University of Tokyo , 7-3-1 Hongo, Bunkyo, Tokyo 113-0033 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Takumu","family":"Kamekawa","sequence":"additional","affiliation":[{"name":"Faculty of Science, Tokyo Metropolitan University , 1-1 Minami-osawa, Hachioji, Tokyo 192-0397 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5109-0024","authenticated-orcid":false,"given":"Takeshi","family":"Izawa","sequence":"additional","affiliation":[{"name":"Graduate School of Agricultural and Life Sciences, The University of Tokyo , 1-1-1 Yayoi, Bunkyo, Tokyo 113-8657 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6531-210X","authenticated-orcid":false,"given":"Takashi","family":"Okamoto","sequence":"additional","affiliation":[{"name":"Graduate School of Science, Tokyo Metropolitan University , 1-1 Minami-osawa, Hachioji, Tokyo 192-0397 ,","place":["Japan"]},{"name":"Faculty of Science, Tokyo Metropolitan University , 1-1 Minami-osawa, Hachioji, Tokyo 192-0397 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,7,31]]},"reference":[{"key":"2026091013263557200_ref1","doi-asserted-by":"publisher","first-page":"1927","DOI":"10.1093\/jxb\/ert054","article-title":"Gene expression profiles in rice gametes and zygotes: identification of gamete-enriched genes and up- or down-regulated genes in zygotes after fertilization","volume":"64","author":"Abiko","year":"2013","journal-title":"J. Exp. Bot."},{"key":"2026091013263557200_ref2","doi-asserted-by":"publisher","first-page":"349","DOI":"10.1016\/j.devcel.2017.10.005","article-title":"The zygotic transition is initiated in unicellular plant zygotes with asymmetric activation of parental genomes","volume":"43","author":"Anderson","year":"2017","journal-title":"Dev. Cell"},{"key":"2026091013263557200_ref3","doi-asserted-by":"publisher","first-page":"1120","DOI":"10.1038\/ncb1201-1120","article-title":"Differential contribution of cytoplasmic Ca2+ and Ca2+ influx to gamete fusion and egg activation in maize","volume":"3","author":"Antoine","year":"2001","journal-title":"Nat. Cell Biol."},{"key":"2026091013263557200_ref4","doi-asserted-by":"publisher","first-page":"2106","DOI":"10.1105\/tpc.17.00099","article-title":"Zygotic genome activation occurs shortly after fertilization in maize","volume":"29","author":"Chen","year":"2017","journal-title":"Plant Cell"},{"key":"2026091013263557200_ref5","doi-asserted-by":"publisher","first-page":"4645","DOI":"10.1038\/ncomms5645","article-title":"Male-female communication triggers calcium signatures during fertilization in Arabidopsis","volume":"5","author":"Denninger","year":"2014","journal-title":"Nat. Commun."},{"key":"2026091013263557200_ref6","doi-asserted-by":"publisher","first-page":"204","DOI":"10.1007\/BF00233634","article-title":"Regeneration and characterization of plants produced from mature tobacco pollen protoplasts via gametosomatic hybridization","volume":"14","author":"Desprez","year":"1995","journal-title":"Plant Cell Rep."},{"key":"2026091013263557200_ref7","doi-asserted-by":"publisher","first-page":"747","DOI":"10.2307\/3869612","article-title":"Karyogamy after electrofusion of single egg and sperm cell protoplasts from maize: cytological evidence and time course","volume":"5","author":"Faure","year":"1993","journal-title":"Plant Cell"},{"key":"2026091013263557200_ref8","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1093\/jxb\/43.3.419","article-title":"A Nicotiana gametosomatic hybrid and its progenies","volume":"43","author":"Giddings","year":"1992","journal-title":"J. Exp. Bot."},{"key":"2026091013263557200_ref9","doi-asserted-by":"publisher","first-page":"4722","DOI":"10.1038\/ncomms5722","article-title":"Live imaging of calcium spikes during double fertilization in Arabidopsis","volume":"5","author":"Hamamura","year":"2014","journal-title":"Nat. Commun."},{"key":"2026091013263557200_ref10","doi-asserted-by":"publisher","first-page":"17159","DOI":"10.1038\/s41598-019-53704-2","article-title":"Transcriptional activation of Arabidopsis zygotes is required for initial cell divisions","volume":"9","author":"Kao","year":"2019","journal-title":"Sci. Rep."},{"key":"2026091013263557200_ref11","doi-asserted-by":"publisher","first-page":"737","DOI":"10.1515\/znc-1973-11-1215","article-title":"The effect of high pH and calcium on tobacco leaf protoplast fusion","volume":"28","author":"Keller","year":"1973","journal-title":"Z. Naturforsch. C"},{"key":"2026091013263557200_ref12","doi-asserted-by":"publisher","first-page":"9652","DOI":"10.1073\/pnas.1821435116","article-title":"DNA demethylation by ROS1a in rice vegetative cells promotes methylation in sperm","volume":"116","author":"Kim","year":"2019","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"2026091013263557200_ref13","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1385\/1-59259-583-9:259","article-title":"In vitro fertilization with isolated single gametes","volume":"111","author":"Kranz","year":"1999","journal-title":"Methods Mol. Biol."},{"key":"2026091013263557200_ref14","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1007\/BF00040086","article-title":"Intra- and interspecific gametosomatic hybridisation within the genus Petunia","volume":"12","author":"Lee","year":"1988","journal-title":"Plant Cell Tissue Organ Cult."},{"key":"2026091013263557200_ref15","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1007\/BF00272968","article-title":"Intraspecific gametosomatic hybridisation in Petunia hybrida","volume":"7","author":"Lee","year":"1988","journal-title":"Plant Cell Rep."},{"key":"2026091013263557200_ref16","first-page":"905","article-title":"Regeneration of hybrid plantlets via pollen-hypocotyl protoplast fusion in Brassica spp","volume":"36","author":"Li","year":"1994","journal-title":"Acta Bot. Sin."},{"key":"2026091013263557200_ref17","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1007\/s00709-006-0228-5","article-title":"Culture and fusion of pollen protoplasts of Brassica oleracea L. var. italica with haploid mesophyll protoplasts of B. rapa L. ssp. pekinensis","volume":"231","author":"Liu","year":"2007","journal-title":"Protoplasma"},{"key":"2026091013263557200_ref18","doi-asserted-by":"publisher","first-page":"6571","DOI":"10.1038\/s41467-023-42394-0","article-title":"Paternal DNA methylation is remodeled to maternal levels in rice zygote","volume":"14","author":"Liu","year":"2023","journal-title":"Nat. Commun."},{"key":"2026091013263557200_ref19","first-page":"342","article-title":"Pollen-mesophyll protoplast fusion and hybrid plant regeneration in Nicotiana","volume":"38","author":"Lu","year":"1996","journal-title":"Acta Bot. Sin."},{"key":"2026091013263557200_ref20","doi-asserted-by":"publisher","first-page":"2369","DOI":"10.1111\/nph.17747","article-title":"Development and regeneration of wheat-rice hybrid zygotes produced by in vitro fertilization system","volume":"232","author":"Maryenti","year":"2021","journal-title":"New Phytol."},{"key":"2026091013263557200_ref21","doi-asserted-by":"publisher","first-page":"1344","DOI":"10.1093\/pcp\/pcae074","article-title":"Wheat cybrid plants, OryzaWheat, regenerated from wheat\u2013rice hybrid zygotes via in vitro fertilization system possess wheat\u2013rice hybrid mitochondria","volume":"65","author":"Maryenti","year":"2024","journal-title":"Plant Cell Physiol."},{"key":"2026091013263557200_ref22","doi-asserted-by":"publisher","first-page":"487","DOI":"10.2307\/3870085","article-title":"Dynamics of nuclear DNA quantities during zygote development in barley","volume":"7","author":"Mogensen","year":"1995","journal-title":"Plant Cell"},{"key":"2026091013263557200_ref23","doi-asserted-by":"publisher","first-page":"641","DOI":"10.1007\/BF00191572","article-title":"Nuclear DNA amounts in the egg and zygote of maize (Zea Mays L.)","volume":"197","author":"Mogensen","year":"1995","journal-title":"Planta"},{"key":"2026091013263557200_ref24","doi-asserted-by":"publisher","first-page":"1747","DOI":"10.1016\/j.febslet.2006.02.028","article-title":"Differential gene expression in egg cells and zygotes suggests that the transcriptome is restructed before the first zygotic division in tobacco","volume":"580","author":"Ning","year":"2006","journal-title":"FEBS Lett."},{"key":"2026091013263557200_ref25","doi-asserted-by":"publisher","first-page":"1533","DOI":"10.1104\/pp.114.236059","article-title":"Dynamics of male and female chromatin during karyogamy in rice zygotes","volume":"165","author":"Ohnishi","year":"2014","journal-title":"Plant Physiol."},{"key":"2026091013263557200_ref26","doi-asserted-by":"publisher","first-page":"5912","DOI":"10.1093\/jxb\/eraf354","article-title":"Inter-subfamily cybrid plants between wheat and maize, Zeawheat: production via an in vitro fertilization system, genome composition, and photosynthetic type","volume":"76","author":"Onda","year":"2025","journal-title":"J. Exp. Bot."},{"key":"2026091013263557200_ref27","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1007\/BF00257851","article-title":"A selection method for the synthesis of triploid hybrids by fusion of microspore protoplasts (n) with somatic cell protoplasts (2n)","volume":"76","author":"Pental","year":"1988","journal-title":"Theor. Appl. Genet."},{"key":"2026091013263557200_ref28","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1007\/BF00261453","article-title":"The production of fertile, triploid somatic hybrid plants (Nicotiana glutinosa (n) + N. tabacum (2n)) via gametic: somatic protoplast fusion","volume":"72","author":"Pirrie","year":"1986","journal-title":"Theor. Appl. Genet."},{"key":"2026091013263557200_ref29","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1093\/pcp\/pcz030","article-title":"Expression of genes from paternal alleles in rice zygotes and involvement of OsASGR-BBML1 in initiation of zygotic development","volume":"60","author":"Rahman","year":"2019","journal-title":"Plant Cell Physiol."},{"key":"2026091013263557200_ref30","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1093\/jxb\/eraf406","article-title":"Autonomous development and regeneration of isolated rice egg cells in vitro in a fertilization-independent manner","volume":"77","author":"Rattanawong","year":"2026","journal-title":"J. Exp. Bot."},{"key":"2026091013263557200_ref31","doi-asserted-by":"publisher","first-page":"377","DOI":"10.5511\/plantbiotechnology.26.377","article-title":"Gametosomatic hybridization between egg cell protoplast and mesophyll protoplast of Petunia hybrida","volume":"26","author":"Sangthong","year":"2009","journal-title":"Plant Biotechnol."},{"key":"2026091013263557200_ref32","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1007\/s00497-009-0129-9","article-title":"Asymmetric cell division of rice zygotes located in embryo sac and produced by in vitro fertilization","volume":"23","author":"Sato","year":"2010","journal-title":"Sex. Plant Reprod."},{"key":"2026091013263557200_ref33","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1007\/s00497-017-0318-x","article-title":"Cell cycle in egg cell and its progression during zygotic development in rice","volume":"31","author":"Sukawa","year":"2018","journal-title":"Plant Reprod."},{"key":"2026091013263557200_ref34","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1104\/pp.15.01953","article-title":"Development of polyspermic rice zygotes","volume":"171","author":"Toda","year":"2016","journal-title":"Plant Physiol."},{"key":"2026091013263557200_ref35","doi-asserted-by":"publisher","DOI":"10.21769\/BioProtoc.2074","article-title":"Electro-fusion of gametes and subsequent culture of zygotes in rice","volume":"6","author":"Toda","year":"2016","journal-title":"Bio-protocol"},{"key":"2026091013263557200_ref36","doi-asserted-by":"publisher","first-page":"2609","DOI":"10.1093\/jxb\/ery094","article-title":"An imbalanced parental genome ratio affects the development of rice zygotes","volume":"69","author":"Toda","year":"2018","journal-title":"J. Exp. Bot."},{"key":"2026091013263557200_ref37","doi-asserted-by":"publisher","first-page":"DEV204497","DOI":"10.1242\/dev.204497","article-title":"Transcriptional dynamics during karyogamy in rice zygotes","volume":"152","author":"Toda","year":"2025","journal-title":"Development"},{"key":"2026091013263557200_ref38","doi-asserted-by":"publisher","first-page":"969","DOI":"10.1111\/j.1365-313X.2006.02836.x","article-title":"Early infection of scutellum tissue with Agrobacterium allows high-speed transformation of rice","volume":"47","author":"Toki","year":"2006","journal-title":"Plant J."},{"key":"2026091013263557200_ref39","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1007\/s00497-006-0020-x","article-title":"Isolation of gametes and central cells from Oryza sativa L","volume":"19","author":"Uchiumi","year":"2006","journal-title":"Sex. Plant Reprod."},{"key":"2026091013263557200_ref40","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1007\/s00425-007-0506-2","article-title":"Establishment of an in vitro fertilization system in rice (Oryza sativa L.)","volume":"226","author":"Uchiumi","year":"2007","journal-title":"Planta"},{"key":"2026091013263557200_ref41","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1111\/j.1365-313X.2010.04403.x","article-title":"Dynamic changes of transcript profiles after fertilization are associated with de novo transcription and maternal elimination in tobacco zygote, and mark the onset of the maternal-to-zygotic transition","volume":"65","author":"Zhao","year":"2011","journal-title":"Plant J."},{"key":"2026091013263557200_ref42","doi-asserted-by":"publisher","first-page":"130","DOI":"10.1104\/pp.16.01406","article-title":"Does early embryogenesis in eudicots and monocots involve the same mechanism and molecular players?","volume":"173","author":"Zhao","year":"2017","journal-title":"Plant Physiol."},{"key":"2026091013263557200_ref43","doi-asserted-by":"publisher","first-page":"882","DOI":"10.1016\/j.devcel.2019.04.016","article-title":"Two-step maternal-to-zygotic transition with two-phase parental genome contributions","volume":"49","author":"Zhao","year":"2019","journal-title":"Dev. Cell"},{"key":"2026091013263557200_ref44","doi-asserted-by":"publisher","first-page":"795","DOI":"10.1038\/s41477-019-0471-3","article-title":"Single-cell three-dimensional genome structures of rice gametes and unicellular zygotes","volume":"5","author":"Zhou","year":"2019","journal-title":"Nat. Plants"}],"container-title":["Plant and Cell Physiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/pcp\/advance-article-pdf\/doi\/10.1093\/pcp\/pcag110\/70412928\/pcag110.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/pcp\/advance-article-pdf\/doi\/10.1093\/pcp\/pcag110\/70412928\/pcag110.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T17:26:46Z","timestamp":1789061206000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/pcp\/advance-article\/doi\/10.1093\/pcp\/pcag110\/8748486"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,31]]},"references-count":44,"URL":"https:\/\/doi.org\/10.1093\/pcp\/pcag110","relation":{},"ISSN":["0032-0781","1471-9053"],"issn-type":[{"value":"0032-0781","type":"print"},{"value":"1471-9053","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,31]]},"article-number":"pcag110"}}