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Research Group for Genomic Epidemiology, Technical University of Denmark, Kgs, Lyngby, DenmarkPatrick Munk, Christian Brinch, Frederik Duus Møller, Thomas N. Petersen, Rene S. Hendriksen, Anne Mette Seyfarth, Jette S. Kjeldgaard, Christina Aaby Svendsen & Frank M. AarestrupCentre for Immunity, Infection and Evolution, University of Edinburgh, Edinburgh, UKBram van Bunnik & Mark WoolhouseCentre for Antibiotic Resistance Research (CARe), University of Gothenburg, Gothenburg, SwedenFanny Berglund & D. G. Joakim LarssonDepartment of Viroscience, Erasmus MC, Rotterdam, The NetherlandsMarion KoopmansInstitute of Public Health, Tirana, AlbaniaArtan BegoUniversidad de Buenos Aires, Buenos Aires, ArgentinaPablo PowerMelbourne Water Corporation, Melbourne, AustraliaCatherine Rees & Kris CoventryCharles Darwin University, Darwin, AustraliaDionisia LambrinidisUniversity of Copenhagen, Frederiksberg C, DenmarkElizabeth Heather Jakobsen Neilson & Yaovi Mahuton Gildas HounmanouCharles Darwin University, Darwin Northern Territory, AustraliaKaren GibbCanberra Hospital, Canberra, AustraliaPeter CollignonALS Water, Scoresby, AustraliaSusan CassarAustrian Agency for Health and Food Safety (AGES), Vienna, AustriaFranz AllerbergerUniversity of Dhaka, Dhaka, BangladeshAnowara Begum & Zenat Zebin HossainEnvironmental Protection Department, Bridgetown, St. Michael, BarbadosCarlon WorrellLaboratoire Hospitalier Universitaire de Bruxelles (LHUB-ULB), Brussels, BelgiumOlivier VandenbergAQUAFIN NV, Aartselaar, BelgiumIlse PietersPolytechnic School of Abomey-Calavi, Abomey-Calavi, BeninDougnon Tamègnon VictorienUniversidad Catσlica Boliviana San Pablo, La Paz, BoliviaAngela Daniela Salazar Gutierrez & Freddy SoriaPublic Health Institute of the Republic of Srpska, Faculty of Medicine University of Banja Luka, Banja Luka, Bosnia and HerzegovinaVesna Rudić GrujićPublic Health Institute of the Republic of Srpska, Banja Luka, Bosnia and HerzegovinaNataša MazalicaBotswana International University of Science and Technology, Palapye, BotswanaTeddie O. RahubeUniversidade Federal de Minas Gerais, Belo Horizonte, BrazilCarlos Alberto Tagliati & Larissa Camila Ribeiro de SouzaOswaldo Cruz Institute, Rio de Janeiro, BrazilDalia RodriguesVale Institute of Technology, Belιm, PA, BrazilGuilherme OliveiraNational Center of Infectious and Parasitic Diseases, Sofia, BulgariaIvan IvanovUniversity of Ouagadougou, Ouagadougou, Burkina FasoBonkoungou Isidore Juste & Traoré OumarInstitut Pasteur du Cambodge, Phnom Penh, CambodiaThet Sopheak & Yith VuthyCentre Pasteur du Cameroun, Yaoundι, CameroonAntoinette Ngandijo, Ariane Nzouankeu & Ziem A. Abah Jacques OlivierUniversity of Regina, Regina, CanadaChristopher K. YostEau Terre Environnement Research Centre (INRS-ETE), Quebec City G1K 9A9, Canada and Indian Institute of Technology, Jammu, IndiaPratik KumarEau Terre Environnement Research Centre (INRS-ETE), Quebec City G1K 9A9, Canada and Lassonde School of Enginerring, York University, Toronto, CanadaSatinder Kaur BrarUniversity of N’Djamena, N’Djamena, ChadDjim-Adjim TaboEscuela de Medicina Veterinaria, Facultad de Ciencias de la Vida, Universidad Andres Bello, Santiago, ChileAiko D. AdellInstitute of Public Health, Santiago, ChileEsteban Paredes-Osses & Maria Cristina MartinezCentro de Biotecnologνa de los Recursos Naturales, Facultad de Ciencias Agrarias y Forestales, Talca, ChileSara Cuadros-OrellanaGuangdong Provincial Center for Disease Control and Prevention, Guangzhou, ChinaChangwen Ke, Huanying Zheng & Li BaishengThe Hong Kong Polytechnic University, Hong Kong, ChinaLok Ting Lau & Teresa ChungShantou University Medical College, Shantou, ChinaXiaoyang JiaoNanjing University of Information Science and Technology, Nanjing, ChinaYongjie YuCenter for Disease Control and Prevention of Henan province, Zhengzhou, ChinaZhao JiaYongColombian Integrated Program for Antimicrobial Resistance Surveillance – Coipars, CI Tibaitatα, Corporaciσn Colombiana de Investigaciσn Agropecuaria (AGROSAVIA), Tibaitatα – Mosquera, Cundinamarca, ColombiaJohan F. Bernal Morales, Maria Fernanda Valencia & Pilar Donado-GodoyInstitut Pasteur de Côte d’Ivoire, Abidjan, Côte d’IvoireKalpy Julien CoulibalyUniversity of Zagreb, Zagreb, CroatiaJasna HrenovicAndrija Stampar Teaching Institute of Public Health, Zagreb, CroatiaMatijana JergovićVeterinary Research Institute, Brno, Czech RepublicRenáta KarpíškováCentre de Recherche en Sciences Naturelles de Lwiro (CRSN-LWIRO), Bukavu, Democratic Republic of CongoZozo Nyarukweba DeogratiasBIOFOS A/S, Copenhagen K, DenmarkBodil ElsborgTechnical University of Denmark, Kgs., Lyngby, DenmarkLisbeth Truelstrup Hansen & Pernille Erland JensenSuez Canal University, Ismailia, EgyptMohamed AbouelnagaUniversity of Sadat City, Sadat City, EgyptMohamed Fathy SalemMinistry of Health, Environmental Microbiology, Tallinn, EstoniaMarliin KoolmeisterAddis Ababa University, Addis Ababa, EthiopiaMengistu Legesse & Tadesse EgualeUniversity of Helsinki, Helsinki, FinlandAnnamari HeikinheimoFrench Institute Search Pour L’exploitation De La Mer (Ifremer), Nantes, FranceSoizick Le Guyader & Julien SchaefferInstituto Nacional de Investigaciσn en Salud Pϊblica-INSPI (CRNRAM), Galαpagos, Quito, EcuadorJose Eduardo VillacisNational Public Health Laboratories, Ministry of Health and Social Welfare, Kotu, GambiaBakary SannehNational Center for Disease Control and Public Health, Tbilisi, GeorgiaLile MalaniaRobert Koch Institute, Berlin, GermanyAndreas Nitsche & Annika BrinkmannTechnische Universitδt Dresden, Institute of Hydrobiology, Dresden, GermanySara Schubert, Sina Hesse & Thomas U. BerendonkUniversity for Development Studies, Tamale, GhanaCourage Kosi Setsoafia SabaUniversity of Ghana, Accra, GhanaJibril MohammedKwame Nkrumah University of Science and Technology, Kumasi, PMB, GhanaPatrick Kwame FegloCouncil for Scientific and Industrial Research Water Research Institute, Accra, GhanaRegina Ama BanuVeterinary Research Institute of Thessaloniki, Hellenic Agricultural Organisation-DEMETER, Thermi, GreeceCharalampos KotzamanidisAthens Water Supply and Sewerage Company (EYDAP S.A.), Athens, GreeceEfthymios LytrasUniversidad de San Carlos de Guatemala, Guatemala City, GuatemalaSergio A. LickesSemmelweis University, Institute of Medical Microbiology, Budapest, HungaryBela KocsisUniversity of Veterinary Medicine, Budapest, HungaryNorbert SolymosiUniversity of Iceland, Reykjavνk, IcelandThorunn R. ThorsteinsdottirCochin University of Science and Technology, Cochin, IndiaAbdulla Mohamed HathaKasturba Medical College, Manipal, IndiaMamatha BallalApollo Diagnostics, Mangalore, IndiaSohan Rodney BangeraShiraz University of Medical Sciences, Shiraz, IranFereshteh FaniShahid Beheshti University of Medical Sciences, Tehran, IranMasoud AlebouyehNational University of Ireland Galway, Galway, IrelandDearbhaile Morris, Louise O’Connor & Martin CormicanBen Gurion University of the Negev and Ministry of Health, Beer-Sheva, IsraelJacob Moran-GiladIstituto Zooprofilattico Sperimentale del Lazio e della Toscana, Rome, ItalyAntonio Battisti, Elena Lavinia Diaconu & Patricia AlbaCNR – Water Research Institute, Verbania, ItalyGianluca Corno & Andrea Di CesareNational Institute of Infectious Diseases, Tokyo, JapanJunzo Hisatsune, Liansheng Yu, Makoto Kuroda, Motoyuki Sugai & Shizuo KayamaNational Center of Expertise, Taldykorgan, KazakhstanZeinegul ShakenovaMount Kenya University, Thika, KenyaCiira KiiyukiaKenya Medical Research Institute, Nairobi, KenyaEric Ng’enoUniversity of Prishtina “Hasan Prishtina” & National Institute of Public Health of Kosovo, Pristina, KosovoLul RakaKuwait Institute for Scientific Research, Kuwait City, KuwaitKazi Jamil, Saja Adel Fakhraldeen & Tareq AlaatiInstitute of Food Safety, Riga, LatviaAivars Bērziņš, Jeļena Avsejenko, Kristina Kokina, Madara Streikisa & Vadims BartkevicsAmerican University of Beirut, Beirut, LebanonGhassan M. MatarCentral Michigan University & Michigan Health Clinics, Saginaw, MI, USAZiad DaoudNational Food and Veterinary Risk Assessment Institute, Vilnius, LithuaniaAsta Pereckienė & Ceslova Butrimaite-AmbrozevicieneLuxembourg Institute of Science and Technology, Belvaux, LuxembourgChristian PennyInstitut Pasteur de Madagascar, Antananarivo, MadagascarAlexandra Bastaraud & Jean-Marc CollardUniversity of Antananarivo, Centre d’Infectiologie Charles Mιrieux, Antananarivo, MadagascarTiavina Rasolofoarison, Luc Hervé Samison & Mala Rakoto AndrianariveloUniversity of Malawi, Blantyre, MalawiDaniel Lawadi BandaMalaysian Genomics Resource Centre Berhad, Kuala Lumpur, MalaysiaArshana AminAIMST University, COMBio, Kedah, MalaysiaHeraa Rajandas & Sivachandran ParimannanWater Services Corporation, Luqa, MaltaDavid SpiteriEnvironmental Health Directorate, St. Venera, MaltaMalcolm Vella HaberUniversity of Mauritius, Reduit, MauritiusSunita J. SantchurnInstitute for Public Health Montenegro, Podgorica, MontenegroAleksandar Vujacic & Dijana DjurovicInstitut Pasteur du Maroc, Casablanca, MoroccoBrahim Bouchrif & Bouchra KarraouanCentro de Investigaηγo em Saϊde de Manhiηa (CISM), Maputo, MozambiqueDelfino Carlos VubilAgriculture and Forestry University, Kathmandu, NepalPushkar PalNational Institute for Public, Health and the Environment (RIVM), Bilthoven, The NetherlandsHeike Schmitt & Mark van PasselUniversity of Otago, Dunedin, New ZealandGert-Jan Jeunen & Neil GemmellUniversity of Otago, Christchurch, New ZealandStephen T. ChambersUniversity of Central America, Managua, NicaraguaFania Perez Mendoza & Jorge Huete-PιrezUniversidad Nacional Autσnoma de Nicaragua-Leσn, Leσn, NicaraguaSamuel VilchezUniversity of Ilorin, Ilorin, NigeriaAkeem Olayiwola Ahmed, Ibrahim Raufu Adisa & Ismail Ayoade OdetokunUniversity of Ibadan, Ibadan, NigeriaKayode FashaeNorwegian Institute of Public Health, Oslo, NorwayAnne-Marie Sørgaard & Astrid Louise WesterVEAS, Slemmestad, NorwayPia Ryrfors & Rune HolmstadUniversity of Agriculture, Faisalabad, PakistanMashkoor MohsinAga Khan University, Karachi, PakistanRumina Hasan & Sadia ShakoorLaboratorio Central de Salud Publica, Asuncion, ParaguayNatalie Weiler Gustafson & Claudia Huber SchillInstituto Nacional de Salud, Lima, PeruMaria Luz Zamudio RojasUniversidad de Piura, Piura, PeruJorge Echevarria Velasquez & Felipe Campos YauceWHO Environmental and Occupational Health, Manila, PhilippinesBonifacio B. MagtibayMaynilad Water Services, Inc., Quezon City, PhilippinesKris Catangcatang & Ruby SibuloNational Veterinary Research Institute, Pulawy, PolandDariusz WasylUniversidade Catσlica Portuguesa, CBQF – Centro de Biotecnologia e Quνmica Fina – Laboratσrio Associado, Escola Superior de Biotecnologia, Porto, PortugalCelia Manaia & Jaqueline RochaAguas do Tejo Atlantico, Lisboa, PortugalJose Martins & Pedro ÁlvaroGwangju Institute of Science and Technology, Gwangju, Republic of KoreaDoris Di Yoong Wen, Hanseob Shin & Hor-Gil HurKorea Advanced Institute of Science and Technology, Daejeon, Republic of KoreaSukhwan YoonInstitute of Public Health of the Republic of North Macedonia, Skopje, Republic of North MacedoniaGolubinka Bosevska & Mihail KochubovskiState Medical and Pharmaceutical University, Chișinău, Republic of MoldovaRadu CojocaruNational Agency for Public Health, Chișinău, Republic of MoldovaOlga BurduniucKing Abdullah University of Science and Technology, Thuwal, Saudi ArabiaPei-Ying HongUniversity of Edinburgh, Edinburgh, Scotland, UKMeghan Rose PerryInstitut Pasteur de Dakar, Dakar, SenegalAmy GassamaInstitute of Veterinary Medicine of Serbia, Belgrade, SerbiaVladimir RadosavljevicNanyang Technological University, Singapore, SingaporeMoon Y. F. Tay, Rogelio Zuniga-Montanez & Stefan WuertzPublic Health Authority of the Slovak Republic, Bratislava, SlovakiaDagmar Gavačová, Katarína Pastuchová & Peter TruskaNational Laboratory of Health, Environment and Food, Ljubljana, SloveniaMarija TrkovIndependent consultant, Johannesburg, South AfricaKaren KeddyDaspoort Waste Water Treatment Works, Pretoria, South AfricaKerneels EsterhuyseKorea Advanced Institute of Science and Technology, Daejeon, South KoreaMin Joon SongSchool of Veterinary Sciences, Lugo, SpainMarcos Quintela-BalujaLabaqua, Santiago de Compostela, SpainMariano Gomez LopezIRTA, Centre de Recerca en Sanitat Animal (CReSA, IRTA-UAB), Campus de la Universitat Autonoma de Barcelona, Bellaterra, SpainMarta Cerdà-CuéllarUniversity of Kelaniya, Ragama, Sri LankaR. R. D. P. Perera, N. K. B. K. R. G. W. Bandara & H. I. PremasiriMedical Research Institute, Colombo, Sri LankaSujatha PathirageCaribbean Public Health Agency, Catries, Saint LuciaKareem CharlemagneThe Sahlgrenska Academy at the University of Gothenburg, Gothenburg, SwedenCarolin RutgerssonSwedish University of Agricultural Sciences, Uppsala, SwedenLeif Norrgren & Stefan ÖrnFederal Food Safety and Veterinary Office (FSVO), Bern, SwitzerlandRenate BossAra Region Bern AG, Herrenschwanden, SwitzerlandTanja Van der HeijdenCenters for Disease Control, Taipei, TaiwanYu-Ping HongKilimanjaro Clinical Research Institute, Moshi, TanzaniaHappiness Houka KumburuSokoine University of Agriculture, Morogoro, TanzaniaRobinson Hammerthon MdegelaFaculty of Science and Technology, Suratthani Rajabhat University, Surat Thani, ThailandKaknokrat ChonsinFaculty of Public Health, Mahidol University, Bangkok, ThailandOrasa SuthienkulFaculty of Medicine Siriraj Hospital, Bangkok, ThailandVisanu ThamlikitkulNational Institute for Public Health and the Environment (RIVM), Bilthoven, NetherlandsAna Maria de Roda HusmanNational Institute of Hygiene, Lomι, TogoBawimodom BidjadaAgence de Mιdecine Prιventive, Dapaong, TogoBerthe-Marie Njanpop-LafourcadeDivision of Integrated Surveillance of Health Emergencies and Response, Lomι, TogoSomtinda Christelle Nikiema-PessinabaPublic Health Institution of Turkey, Ankara, TurkeyBelkis LeventHatay Mustafa Kemal University, Hatay, TurkeyCemil KurekciMakerere University, Kampala, UgandaFrancis Ejobi & John Bosco KaluleAbu Dhabi Public Health Center, Abu Dhai, United Arab EmiratesJens ThomsenDubai municipality, WWTP Al Aweer, Dubai, UAEOuidiane ObaidiRashid Hospital, Dubai, UAELaila Mohamed JassimNorthumbrian Water, Northumbria House, Abbey Road, Pity Me, Durham, UKAndrew MooreUniversity of Exeter Medical School, Cornwall, UKAnne Leonard, Lihong Zhang & William H. GazeNewcastle University, Newcastle upon Tyne, UKDavid W. Graham & Joshua T. BunceBrightwater Treatment Plant, Woodinville, WA, USABrett LeforDepartment of Environmental Sciences and Engineering, University of North Carolina at Chapel Hill, Chapel Hill, NC, USADrew Capone & Joe BrownUniversity of North Carolina, Chapel Hill, USAEmanuele Sozzi & Mark D. SobseyUniversity of Washington, Seattle, WA, USAJohn Scott Meschke, Nicola Koren Beck, Pardi Sukapanpatharam & Phuong TruongBaylor University, Waco, USAMichael DavisColumbia Boulevard WWTP, Portland, USARonald LilienthalEastern Illinois University, Charleston, USASanghoon KangThe Ohio State University, Columbus Ohio, USAThomas E. WittumLaboratorio Tecnolσgico del Uruguay, Montevideo, UruguayNatalia Rigamonti & Patricia BaklayanInstitute of Public Health in Ho Chi Minh City, Ho Chi Minh, VietnamChinh Dang Van, Doan Minh Nguyen Tran & Nguyen Do PhucUniversity of Zambia, Lusaka, ZambiaGeoffrey KwendaF.M.A., M.K., and M.W. conceived the study and secured funding. R.S.H., A.M.S., C.A.A.S., and J.S.K. organized sample collection, material transfer, and logistics. F.D.M., P.M., and C.B. did quality control, sample selection, and outlier detection. P.M., C.B., F.D.M., T.N.P., and F.B. performed bioinformatics analyses. P.M. and C.B. carried out data and statistical analyses and visualization. P.M. and F.M.A. drafted the initial manuscript with input from C.B., B.v.B., D.G.J.L., M.W., and M.K. The Global Sewage Consortium authors carried out sewage sampling, filled in metadata and shipped the samples to DTU. All authors helped to review and improve the manuscript. More

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Time-course observations on 43 flowers of Causonis japonica revealed changes in flower disc colour and sexual expression (Table 1). Temporal changes in floral features showed no difference between diploid (19 flowers) and triploid (24 flowers) individuals. For example, flowering onset times did not differ substantially between ploidy level (diploid: from 07:07 to 13:27, triploid: from 06:58 to 14:49). However, the flowering duration varied significantly from flower to flower, ranging from a minimum of one day to a maximum of six days. Regardless of the ploidy level, all flowers with damaged styles (14 flowers) exhibited brown stigmas after the male phase, then ceased floral development prior to the female phase.Table 1 Characteristics of 43 flowers of Causonis japonica.Full size tableFigure 1 shows the typical time-course changes of C. japonica flower features (flower ID 35 in Table 1) according to the RGB values (representing the activities of nectar secretion: see below). As in the other 42 examined cases shown in Table 1, the initial colour of this flower disc immediately after anthesis (male phase) was orange (Stage 1, Fig. 1a, RGB: 255, 88, 16), as reported earlier7. Immediately after the petals and stamens fell off, the flower disc colour changed to pink (Stage 2, Fig. 1b, RGB: 255, 82, 102). The styles were not yet elongated at this stage, and the flowers were asexual. In 12 cases with damaged styles and brown stigmas, the flower discs remained pink until the flowers fell off (shown as “O–P” in Table 1).Figure 1Colour change process of a flower disc in Causonis japonica (flower ID 35 in Table 1). Disc colour changed from orange (a, RGB: 255, 88, 16) to pink (b, RGB: 255, 82, 102) before recovering to orange (c, RGB: 255, 88, 16) again, then pink (d, RGB: 255, 120, 94) again. In the last stage, the flower disc turned brownish pink (e, RGB: 232, 162, 169) then fell off. The two orange colour stages were synchronised with flower sexual activity. (a) First orange stage shows stamen activity (male phase); (c) second orange stage indicates stigma maturation (female phase). Nectar secretion was active only in the orange stages and more active in the female phase; the same tendency was observed in the other cases shown in Table 1.Full size imageHowever, in the remaining 31 flowers with normally elongated styles, maturation of the styles (female phase) coincided with the flower discs again exhibiting a distinct orange colour (Stage 3, Fig. 1c, RGB: 255, 88, 16). After the female phase, the flower discs turned pink again (Stage 4, shown as “O–P–O–P” in Table 1), and brownish colouration appeared in the stigmas (Fig. 1d, RGB: 255, 120, 94). Finally, the flower discs turned to a faded pink (Fig. 1e, RGB: 232, 162, 169) just before the flowers fell off. Therefore, the above observations imply that colour-change has a strict correlation with sexual phase.The timings of the disc colour change to the second orange stage (female phase) varied depending on the onset time of each flower. Most flowers that opened before 10:00 reached the second orange stage (female phase) on the afternoon of the same day (except for two flowers, ID 1 and 2 in Table 1). Conversely, flowers that bloomed after 10:00 reached the second orange stage (female phase) at approximately noon the following day. These flowering processes were not fully synchronised in the same inflorescence; therefore, pink and orange discs often coexisted in the same inflorescence. Indeed, we can collect various stages of flowers at a time point from one population as shown in Fig. 2a.Figure 2Histology of floral discs of C. japonica. (a) Floral disc colour change observed in a triploid individual. Flowers were hand-sectioned along the longitudinal axis to show inside colouration of floral disc. Floral phase was judged from the stigma length and colour of the stigma tip; from left, initial stage with orange floral disc and short style, first pink stage with short style, second orange stage with elongated style with matured stigma, and second orange stage with elongated style. Unit of scale bar = 1 mm. (b–e) Longitudinal sections of floral discs in the initial orange stage (b, d) and pink stage (c, e). Scale bar = 500 µm. (b, c) Hand sections of living floral discs showing pigmentation of vacuoles in some scattered cells. (d, e) Resin-embedded sections of floral discs showing histology.Full size imageFigure 1 also shows the typical time-course changes of nectar activities (flower ID 35 in Table 1), which indicates active nectar secretions during both orange colour stages. That is, the flower discs secreted nectar in both male and female phases, with no visible nectar secretion in the pink stages. Moreover, nectar secretion in the female phase of this flower was higher than that in the male phase, a tendency that was also observed in other flowers; however, the total volume of nectar varied among the flowers shown in Table 1. During anthesis, we confirmed that bees, wasps, ants and other insects visited the flowers as previously described7 (Supplementary Fig. 1).Longitudinal sections of flowers in the pink-coloured and orange-coloured stages (Fig. 2a) revealed that pigmentation occurred only in a subset of parenchymatous cells in both cases (Fig. 2b, c). No structural or cytological changes were observed between the initial orange stage and the pink stages (Fig. 2d, e), suggesting that the observed oscillating colour change depends on the degradation and biosynthesis of orange pigments.To understand what pigments are involved in the dual colour change of the C. japonica flower disc, we extracted carotenoids and chlorophyll with Acetone. Anthocyanin was also extracted with Methanol-HCl. As a result, while anthocyanin content was not significantly altered throughout the stages examined, we found that carotenoid level strongly correlated with the colour change detected by naked eye. More specifically, in stages 1 and 3 the carotenoid content was high (63.8 and 65.3 µg/g dry weight, respectively), but significantly decreased in stages 2 and 4 (14.3 and 36.5 µg/g dry weight, respectively) (Table 2). Interestingly, an increase in chlorophyll content was confined to stage 4 (Table 2). Together, the observed dual colour change was ascribed to the decrease (stage 2) and the increase (stage 3) of carotenoid contents in the flower discs.Table 2 Chlorophyll and Carotenoid contents in the flower discs of C. japonica.Full size table More

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