{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,21]],"date-time":"2026-04-21T18:11:18Z","timestamp":1776795078344,"version":"3.51.2"},"reference-count":89,"publisher":"The Royal Society","issue":"2011","license":[{"start":{"date-parts":[[2023,11,29]],"date-time":"2023-11-29T00:00:00Z","timestamp":1701216000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2023,11,29]],"date-time":"2023-11-29T00:00:00Z","timestamp":1701216000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100006390","name":"Universit\u00e9 de Lausanne","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100006390","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001711","name":"Swiss National Science Foundation","doi-asserted-by":"crossref","award":["SNSF grant 310030_185196"],"award-info":[{"award-number":["SNSF grant 310030_185196"]}],"id":[{"id":"10.13039\/501100001711","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["royalsocietypublishing.org"],"crossmark-restriction":true},"short-container-title":["Proc. R. Soc. B."],"published-print":{"date-parts":[[2023,11,29]]},"abstract":"<jats:p>\n            In dioecious populations, males and females may evolve different trait values to increase fitness through their respective sexual functions. Because hermaphrodites express both sexual functions, resolving sexual conflict is potentially more difficult for them. Here, we show that hermaphrodite plants can partially resolve sexual conflict by expressing different trait values in different male and female modules (e.g. different flowers, inflorescences, branches etc.). We analysed the flowering phenology, sex allocation and selection gradients on floral traits of flowers of the andromonoecious plant\n            <jats:italic>Pulsatilla alpina<\/jats:italic>\n            , which produces both bisexual and male flowers. Our results indicate that strong protogyny prevents early bisexual flowers from profiting from high siring opportunities early in the reproductive season at a time when male flowers could achieve high siring success. The production of unisexual male flowers thus resolves this sexual conflict because it allows the flowers to express their male function without waiting until after the female function has been performed. Our study illustrates the resolution of sexual conflict arising from phenological constraints via modular divergence in sex allocation. We discuss the extent to which modular variation in sex allocation in the context of other sexual systems may be similarly explained.\n          <\/jats:p>","DOI":"10.1098\/rspb.2023.2137","type":"journal-article","created":{"date-parts":[[2023,11,29]],"date-time":"2023-11-29T08:11:33Z","timestamp":1701245493000},"update-policy":"https:\/\/doi.org\/10.1098\/crossmark-policy","source":"Crossref","is-referenced-by-count":11,"title":["Unisexual flowers as a resolution to intralocus sexual conflict in hermaphrodites"],"prefix":"10.1098","volume":"290","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8056-5858","authenticated-orcid":true,"given":"Kai-Hsiu","family":"Chen","sequence":"first","affiliation":[{"name":"Department of Ecology and Evolution, University of Lausanne, Biophore Building, 1015 Lausanne, Switzerland"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0098-7074","authenticated-orcid":true,"given":"John R.","family":"Pannell","sequence":"additional","affiliation":[{"name":"Department of Ecology and Evolution, University of Lausanne, Biophore Building, 1015 Lausanne, Switzerland"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"175","published-online":{"date-parts":[[2023,11,29]]},"reference":[{"key":"e_1_3_8_2_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.0014-3820.2000.tb00005.x"},{"key":"e_1_3_8_3_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0169-5347(02)02569-7"},{"key":"e_1_3_8_4_2","doi-asserted-by":"publisher","DOI":"10.1038\/nrg.2017.83"},{"key":"e_1_3_8_5_2","doi-asserted-by":"publisher","DOI":"10.2307\/2407393"},{"key":"e_1_3_8_6_2","doi-asserted-by":"publisher","DOI":"10.2307\/2408385"},{"key":"e_1_3_8_7_2","doi-asserted-by":"publisher","DOI":"10.1086\/285655"},{"key":"e_1_3_8_8_2","doi-asserted-by":"publisher","DOI":"10.1111\/evo.12536"},{"key":"e_1_3_8_9_2","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2010.1401"},{"key":"e_1_3_8_10_2","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a017673"},{"key":"e_1_3_8_11_2","doi-asserted-by":"publisher","DOI":"10.1093\/icb\/icj038"},{"key":"e_1_3_8_12_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2435.2006.01179.x"},{"key":"e_1_3_8_13_2","doi-asserted-by":"publisher","DOI":"10.1890\/0012-9658(2000)081[0532:COAAGT]2.0.CO;2"},{"key":"e_1_3_8_14_2","doi-asserted-by":"publisher","DOI":"10.1093\/aob\/mcr193"},{"key":"e_1_3_8_15_2","doi-asserted-by":"publisher","DOI":"10.1038\/sj.hdy.6800402"},{"key":"e_1_3_8_16_2","doi-asserted-by":"publisher","DOI":"10.1038\/sj.hdy.6800020"},{"key":"e_1_3_8_17_2","doi-asserted-by":"publisher","DOI":"10.3732\/ajb.0800106"},{"key":"e_1_3_8_18_2","doi-asserted-by":"publisher","DOI":"10.2307\/2410293"},{"key":"e_1_3_8_19_2","first-page":"1131","article-title":"Temporal floral sex allocation in protogynous Aquilegia yabeana contrasts with protandrous species: support for the mating environment hypothesis","volume":"58","author":"Huang S-Q","year":"2004","unstructured":"Huang S-Q, Tang L-L, Yu Q, Guo Y-H. 2004 Temporal floral sex allocation in protogynous Aquilegia yabeana contrasts with protandrous species: support for the mating environment hypothesis. Evolution 58, 1131-1134. (doi:10.1111\/j.0014-3820.2004.tb00446.x)","journal-title":"Evolution"},{"key":"e_1_3_8_20_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1558-5646.2008.00394.x"},{"key":"e_1_3_8_21_2","doi-asserted-by":"publisher","DOI":"10.1038\/35068635"},{"key":"e_1_3_8_22_2","doi-asserted-by":"publisher","DOI":"10.1080\/0028825X.1986.10409725"},{"key":"e_1_3_8_23_2","doi-asserted-by":"publisher","DOI":"10.3732\/ajb.1400196"},{"key":"e_1_3_8_24_2","doi-asserted-by":"publisher","DOI":"10.1086\/378659"},{"key":"e_1_3_8_25_2","doi-asserted-by":"publisher","DOI":"10.1093\/aob\/mcr025"},{"key":"e_1_3_8_26_2","doi-asserted-by":"publisher","DOI":"10.1093\/aob\/mcp035"},{"key":"e_1_3_8_27_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.0014-3820.2002.tb00847.x"},{"key":"e_1_3_8_28_2","doi-asserted-by":"publisher","DOI":"10.2307\/2408842"},{"key":"e_1_3_8_29_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1461-0248.2011.01601.x"},{"key":"e_1_3_8_30_2","doi-asserted-by":"publisher","DOI":"10.1111\/evo.13639"},{"key":"e_1_3_8_31_2","doi-asserted-by":"publisher","DOI":"10.1086\/376574"},{"key":"e_1_3_8_32_2","first-page":"1458","article-title":"Components of phenotypic selection: pollen export and flower corolla width in Ipomopsis aggregata","volume":"45","author":"Campbell DR","year":"1991","unstructured":"Campbell DR, Waser NM, Price MV, Lynch EA, Mitchell RJ. 1991 Components of phenotypic selection: pollen export and flower corolla width in Ipomopsis aggregata. Evolution 45, 1458-1467. (doi:10.1111\/J.1558-5646.1991.TB02648.X)","journal-title":"Evolution"},{"key":"e_1_3_8_33_2","doi-asserted-by":"publisher","DOI":"10.1086\/316997"},{"key":"e_1_3_8_34_2","first-page":"112","article-title":"Phenotypic selection in hawkmoth-pollinated Plantathera bifolia: targets and fitness surfaces","volume":"54","author":"Maad J","year":"2000","unstructured":"Maad J. 2000 Phenotypic selection in hawkmoth-pollinated Plantathera bifolia: targets and fitness surfaces. Evolution 54, 112-123. (doi:10.1111\/j.0014-3820.2000.tb00012.x)","journal-title":"Evolution"},{"key":"e_1_3_8_35_2","volume-title":"Flora helvetica","author":"Lauber K","year":"2018","unstructured":"Lauber K, Wagner G, Gygax A, Eggenberg S, Michel A. 2018 Flora helvetica, 5th edn. Bern, Switzerland: Haupt Verlag.","edition":"5"},{"key":"e_1_3_8_36_2","doi-asserted-by":"crossref","unstructured":"Chen K-H Pannell JR. 2023 Size-dependent sex allocation and the expression of andromonoecy in a protogynous perennial herb: both size and timing matter. bioRxiv 2023.03.10.532080. (doi:10.1101\/2023.03.10.532080)","DOI":"10.1101\/2023.03.10.532080"},{"key":"e_1_3_8_37_2","doi-asserted-by":"publisher","DOI":"10.1002\/ajb2.16073"},{"key":"e_1_3_8_38_2","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2022.0919"},{"key":"e_1_3_8_39_2","doi-asserted-by":"publisher","DOI":"10.2307\/2409284"},{"key":"e_1_3_8_40_2","volume-title":"R: a language and environment for statistical computing","author":"R Core Team","year":"2021","unstructured":"R Core Team. 2021 R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing."},{"key":"e_1_3_8_41_2","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v067.i01"},{"key":"e_1_3_8_42_2","unstructured":"Hartig F. 2019 DHARMa: residual diagnostics for hierarchical (multi-level\/mixed) regression models. R package version 0.2.4. See https:\/\/cran.r-project.org\/web\/packages\/DHARMa\/vignettes\/DHARMa.html."},{"key":"e_1_3_8_43_2","unstructured":"Pinheiro J Bates D DebRoy S Sarkar D R Core Team. 2022 nlme: linear and nonlinear mixed effects models. See https:\/\/cran.r-project.org\/web\/packages\/nlme\/nlme.pdf."},{"key":"e_1_3_8_44_2","unstructured":"Lenth RV. 2020 Emmeans: estimated marginal means aka least-squares means. See https:\/\/cran.r-project.org\/web\/packages\/emmeans\/index.html."},{"key":"e_1_3_8_45_2","doi-asserted-by":"publisher","DOI":"10.1086\/595841"},{"key":"e_1_3_8_46_2","doi-asserted-by":"publisher","DOI":"10.1515\/9781400850600"},{"key":"e_1_3_8_47_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1558-5646.2010.01021.x"},{"key":"e_1_3_8_48_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1558-5646.2008.00449.x"},{"key":"e_1_3_8_49_2","doi-asserted-by":"publisher","DOI":"10.1111\/evo.14113"},{"key":"e_1_3_8_50_2","doi-asserted-by":"publisher","DOI":"10.3732\/ajb.89.4.587"},{"key":"e_1_3_8_51_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1756-1051.2010.00798.x"},{"key":"e_1_3_8_52_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1420-9101.2005.00896.x"},{"key":"e_1_3_8_53_2","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1093\/acprof:oso\/9780199208784.003.0021","volume-title":"Sex, size, and gender roles: evolutionary studies of sexual size dimorphism","author":"Bedhomme S","year":"2007","unstructured":"Bedhomme S, Chippindale AK. 2007 Irreconcilable differences: when sexual dimorphism fails to resolve sexual conflict. In Sex, size, and gender roles: evolutionary studies of sexual size dimorphism (eds DJ Fairbairn, WU Blanckenhorn, T Sz\u00e9kely), pp. 185-194. New York, NY: Oxford University Press."},{"key":"e_1_3_8_54_2","doi-asserted-by":"publisher","DOI":"10.1007\/BF02857918"},{"key":"e_1_3_8_55_2","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-7091-6670-3"},{"key":"e_1_3_8_56_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ecolsys.33.010802.150419"},{"key":"e_1_3_8_57_2","doi-asserted-by":"publisher","DOI":"10.1007\/BF01508307"},{"key":"e_1_3_8_58_2","doi-asserted-by":"publisher","DOI":"10.1046\/j.1420-9101.1999.00001.x"},{"key":"e_1_3_8_59_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0169-5347(00)01872-3"},{"key":"e_1_3_8_60_2","doi-asserted-by":"publisher","DOI":"10.2307\/2389709"},{"key":"e_1_3_8_61_2","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2001.1771"},{"key":"e_1_3_8_62_2","doi-asserted-by":"publisher","DOI":"10.1038\/263125a0"},{"key":"e_1_3_8_63_2","doi-asserted-by":"publisher","DOI":"10.1890\/03-0522"},{"key":"e_1_3_8_64_2","first-page":"1201","article-title":"Effects of sib-competition on female reproductive success in Salvia lutescens Koidz. var. crenata","volume":"7","author":"Takahashi K","year":"2005","unstructured":"Takahashi K, Makino TT, Sakai S. 2005 Effects of sib-competition on female reproductive success in Salvia lutescens Koidz. var. crenata. Evol. Ecol. Res. 7, 1201-1212.","journal-title":"Evol. Ecol. Res."},{"key":"e_1_3_8_65_2","doi-asserted-by":"publisher","DOI":"10.1890\/0012-9658(1998)079[2771:MDOSDI]2.0.CO;2"},{"key":"e_1_3_8_66_2","doi-asserted-by":"publisher","DOI":"10.1093\/aob\/mcl292"},{"key":"e_1_3_8_67_2","first-page":"139","volume-title":"Plant reproductive ecology: patterns and strategies","author":"Schlessman MA","year":"1988","unstructured":"Schlessman MA. 1988 Gender diphasy (sex choice). In Plant reproductive ecology: patterns and strategies (eds J Lovett-Doust, L Lovett-Doust), pp. 139-153. New York, NY: Oxford University Press."},{"key":"e_1_3_8_68_2","doi-asserted-by":"publisher","DOI":"10.1046\/j.1442-1984.2001.00058.x"},{"key":"e_1_3_8_69_2","doi-asserted-by":"publisher","DOI":"10.1086\/689616"},{"key":"e_1_3_8_70_2","doi-asserted-by":"publisher","DOI":"10.1111\/jeb.12333"},{"key":"e_1_3_8_71_2","doi-asserted-by":"publisher","DOI":"10.1086\/406066"},{"key":"e_1_3_8_72_2","doi-asserted-by":"publisher","DOI":"10.1086\/282974"},{"key":"e_1_3_8_73_2","doi-asserted-by":"publisher","DOI":"10.1159\/000223079"},{"key":"e_1_3_8_74_2","first-page":"128","article-title":"Mating behavior and hermaphroditism in coral reef fishes","volume":"72","author":"Warner RR","year":"1984","unstructured":"Warner RR. 1984 Mating behavior and hermaphroditism in coral reef fishes. Am. Sci. 72, 128-136.","journal-title":"Am. Sci."},{"key":"e_1_3_8_75_2","doi-asserted-by":"publisher","DOI":"10.1086\/283342"},{"key":"e_1_3_8_76_2","first-page":"1087","article-title":"Sex allocation in plants and the evolution of monoecy","volume":"10","author":"de Jong TJ","year":"2008","unstructured":"de Jong TJ, Shmida A, Thuijsman F. 2008 Sex allocation in plants and the evolution of monoecy. Evol. Ecol. Res. 10, 1087-1109.","journal-title":"Evol. Ecol. Res."},{"key":"e_1_3_8_77_2","doi-asserted-by":"publisher","DOI":"10.1080\/0028825X.1986.10409726"},{"key":"e_1_3_8_78_2","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2003.1301"},{"key":"e_1_3_8_79_2","doi-asserted-by":"publisher","DOI":"10.1002\/ajb2.16209"},{"key":"e_1_3_8_80_2","doi-asserted-by":"publisher","DOI":"10.1093\/aobpla\/plv008"},{"key":"e_1_3_8_81_2","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2000.1395"},{"key":"e_1_3_8_82_2","doi-asserted-by":"publisher","DOI":"10.1111\/mec.16820"},{"key":"e_1_3_8_83_2","doi-asserted-by":"publisher","DOI":"10.1086\/712646"},{"key":"e_1_3_8_84_2","doi-asserted-by":"crossref","unstructured":"Larue C Petit RJ. 2023 Harmful self-pollination drives gynodioecy in European chestnut a self-incompatible tree. bioRxiv 2022.08.01.502348. (doi:10.1101\/2022.08.01.502348)","DOI":"10.1101\/2022.08.01.502348"},{"key":"e_1_3_8_85_2","doi-asserted-by":"publisher","DOI":"10.1007\/s00606-003-0029-7"},{"key":"e_1_3_8_86_2","doi-asserted-by":"publisher","DOI":"10.2307\/2446311"},{"key":"e_1_3_8_87_2","doi-asserted-by":"publisher","DOI":"10.3724\/SP.J.1258.2014.00008"},{"key":"e_1_3_8_88_2","doi-asserted-by":"publisher","DOI":"10.1111\/evo.13572"},{"key":"e_1_3_8_89_2","doi-asserted-by":"crossref","unstructured":"Chen K-H Pannell JR. 2023 Data from: Unisexual flowers as a resolution to intralocus sexual conflict in hermaphrodites. Dryad Digital Repository. (doi:10.5061\/dryad.rbnzs7hj0)","DOI":"10.1098\/rspb.2023.2137"},{"key":"e_1_3_8_90_2","doi-asserted-by":"crossref","unstructured":"Chen K-H Pannell JR. 2023 Data from: Unisexual flowers as a resolution to intralocus sexual conflict in hermaphrodites. Figshare. (doi:10.6084\/m9.figshare.c.6927463)","DOI":"10.1098\/rspb.2023.2137"}],"container-title":["Proceedings of the Royal Society B: Biological Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/doi\/pdf\/10.1098\/rspb.2023.2137","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/royalsocietypublishing.org\/doi\/full-xml\/10.1098\/rspb.2023.2137","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/royalsocietypublishing.org\/doi\/pdf\/10.1098\/rspb.2023.2137","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,1,9]],"date-time":"2024-01-09T09:41:47Z","timestamp":1704793307000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/doi\/10.1098\/rspb.2023.2137"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,11,29]]},"references-count":89,"journal-issue":{"issue":"2011","published-print":{"date-parts":[[2023,11,29]]}},"alternative-id":["10.1098\/rspb.2023.2137"],"URL":"https:\/\/doi.org\/10.1098\/rspb.2023.2137","relation":{},"ISSN":["0962-8452","1471-2954"],"issn-type":[{"value":"0962-8452","type":"print"},{"value":"1471-2954","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,11,29]]},"assertion":[{"value":"2023-09-19","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2023-11-01","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2023-11-29","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"20232137"}}