{"id":"https://openalex.org/W7214099504","doi":"https://doi.org/10.64898/2026.09.17.752332","title":"Phenotype-dependent schooling behaviour across hierarchical levels in three-spined sticklebacks","display_name":"Phenotype-dependent schooling behaviour across hierarchical levels in three-spined sticklebacks","publication_year":2026,"publication_date":"2026-09-23","ids":{"openalex":"https://openalex.org/W7214099504","doi":"https://doi.org/10.64898/2026.09.17.752332"},"language":"en","primary_location":{"id":"doi:10.64898/2026.09.17.752332","doi":"https://doi.org/10.64898/2026.09.17.752332","is_oa":true,"landing_page_url":"https://doi.org/10.64898/2026.09.17.752332","pdf_url":"https://www.biorxiv.org/content/biorxiv/early/2026/09/23/2026.09.17.752332.full.pdf","source":{"id":"https://openalex.org/S4306402567","display_name":"bioRxiv (Cold Spring Harbor Laboratory)","issn_l":null,"issn":null,"is_oa":false,"is_in_doaj":false,"is_core":false,"listed_in":[],"host_organization":"https://openalex.org/I2750212522","host_organization_name":"Cold Spring Harbor Laboratory","host_organization_lineage":["https://openalex.org/I2750212522"],"host_organization_lineage_names":[],"type":"repository"},"license":"cc-by-nc","license_id":"https://openalex.org/licenses/cc-by-nc","version":"acceptedVersion","is_accepted":true,"is_published":false,"raw_source_name":"bioRxiv","raw_type":"posted-content"},"volume":null,"type":"preprint","indexed_in":["crossref"],"open_access":{"is_oa":true,"oa_status":"green","oa_url":"https://www.biorxiv.org/content/biorxiv/early/2026/09/23/2026.09.17.752332.full.pdf","any_repository_has_fulltext":true},"authorships":[{"author_position":"first","author":{"id":"https://openalex.org/A5001311794","display_name":"Aparajitha Ramesh","orcid":"https://orcid.org/0000-0002-7200-1366","observed_orcids":["https://orcid.org/0000-0002-7200-1366"]},"institutions":[{"id":"https://openalex.org/I118564535","display_name":"University of Bern","ror":"https://ror.org/02k7v4d05","country_code":"CH","type":"education","lineage":["https://openalex.org/I118564535"]}],"countries":["CH"],"is_corresponding":false,"raw_author_name":"Aparajitha Ramesh","raw_affiliation_strings":["Behavioural Ecology, Institute of Ecology and Evolution, University of Bern, Wohlenstrasse 50A, CH-3032 Hinterkappelen, Switzerland"],"raw_orcid":"https://orcid.org/0000-0002-7200-1366","affiliations":[{"raw_affiliation_string":"Behavioural Ecology, Institute of Ecology and Evolution, University of Bern, Wohlenstrasse 50A, CH-3032 Hinterkappelen, Switzerland","institution_ids":["https://openalex.org/I118564535"]}]},{"author_position":"middle","author":{"id":"https://openalex.org/A5046364260","display_name":"Eize J. Stamhuis","orcid":"https://orcid.org/0000-0001-7746-2535","observed_orcids":["https://orcid.org/0000-0001-7746-2535"]},"institutions":[{"id":"https://openalex.org/I169381384","display_name":"University of Groningen","ror":"https://ror.org/012p63287","country_code":"NL","type":"education","lineage":["https://openalex.org/I169381384"]}],"countries":["NL"],"is_corresponding":false,"raw_author_name":"Eize Stamhuis","raw_affiliation_strings":["Department of Oceans Ecosystems, Energy and Sustainability Research Institute Groningen, University of Groningen, Nijenborgh 7, 9747 AG Groningen, Netherlands"],"raw_orcid":"https://orcid.org/0000-0001-7746-2535","affiliations":[{"raw_affiliation_string":"Department of Oceans Ecosystems, Energy and Sustainability Research Institute Groningen, University of Groningen, Nijenborgh 7, 9747 AG Groningen, Netherlands","institution_ids":["https://openalex.org/I169381384"]}]},{"author_position":"last","author":{"id":"https://openalex.org/A5042660363","display_name":"Marion Nicolaus","orcid":"https://orcid.org/0000-0003-1808-1526","observed_orcids":["https://orcid.org/0000-0003-1808-1526"]},"institutions":[{"id":"https://openalex.org/I169381384","display_name":"University of Groningen","ror":"https://ror.org/012p63287","country_code":"NL","type":"education","lineage":["https://openalex.org/I169381384"]}],"countries":["NL"],"is_corresponding":true,"raw_author_name":"Marion Nicolaus","raw_affiliation_strings":["Conservation Ecology group, Groningen Institute for Evolutionary Life Sciences, University of Groningen, Nijenborgh 7, 9747 AG Groningen, Netherlands"],"raw_orcid":"https://orcid.org/0000-0003-1808-1526","affiliations":[{"raw_affiliation_string":"Conservation Ecology group, Groningen Institute for Evolutionary Life Sciences, University of Groningen, Nijenborgh 7, 9747 AG Groningen, Netherlands","institution_ids":["https://openalex.org/I169381384"]}]}],"institutions":[],"countries_distinct_count":2,"institutions_distinct_count":2,"corresponding_author_ids":["https://openalex.org/A5042660363"],"corresponding_institution_ids":["https://openalex.org/I169381384"],"apc_list":null,"apc_paid":null,"fwci":null,"has_fulltext":true,"cited_by_count":0,"citation_normalized_percentile":null,"cited_by_percentile_year":null,"biblio":{"volume":null,"issue":null,"first_page":null,"last_page":null},"is_retracted":false,"is_paratext":false,"is_xpac":false,"primary_topic":{"id":"https://openalex.org/T10174","display_name":"Animal Behavior and Reproduction","score":0.9609000086784363,"subfield":{"id":"https://openalex.org/subfields/1105","display_name":"Ecology, Evolution, Behavior and Systematics"},"field":{"id":"https://openalex.org/fields/11","display_name":"Agricultural and Biological Sciences"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}},"topics":[{"id":"https://openalex.org/T10174","display_name":"Animal Behavior and Reproduction","score":0.9609000086784363,"subfield":{"id":"https://openalex.org/subfields/1105","display_name":"Ecology, Evolution, Behavior and Systematics"},"field":{"id":"https://openalex.org/fields/11","display_name":"Agricultural and Biological Sciences"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}},{"id":"https://openalex.org/T12014","display_name":"Fish biology, ecology, and behavior","score":0.015799999237060547,"subfield":{"id":"https://openalex.org/subfields/2309","display_name":"Nature and Landscape Conservation"},"field":{"id":"https://openalex.org/fields/23","display_name":"Environmental Science"},"domain":{"id":"https://openalex.org/domains/3","display_name":"Physical Sciences"}},{"id":"https://openalex.org/T10249","display_name":"Distributed Control Multi-Agent Systems","score":0.00570000009611249,"subfield":{"id":"https://openalex.org/subfields/1705","display_name":"Computer Networks and Communications"},"field":{"id":"https://openalex.org/fields/17","display_name":"Computer Science"},"domain":{"id":"https://openalex.org/domains/3","display_name":"Physical Sciences"}}],"keywords":[{"id":"https://openalex.org/keywords/schooling-behavior","display_name":"schooling behavior","score":0.9789999723434448},{"id":"https://openalex.org/keywords/gasterosteus-aculeatus","display_name":"Gasterosteus aculeatus","score":0.9990000128746033},{"id":"https://openalex.org/keywords/anadromous-migration","display_name":"anadromous migration","score":0.7039999961853027},{"id":"https://openalex.org/keywords/landlocked-populations","display_name":"landlocked populations","score":0.8140000104904175},{"id":"https://openalex.org/keywords/collective-behavior","display_name":"collective behavior","score":0.7170000076293945},{"id":"https://openalex.org/keywords/self-organization","display_name":"self-organization","score":0.7379999756813049},{"id":"https://openalex.org/keywords/group-alignment","display_name":"group alignment","score":0.6159999966621399},{"id":"https://openalex.org/keywords/behavioral-plasticity","display_name":"behavioral plasticity","score":0.49000000953674316}],"concepts":[{"id":"https://openalex.org/C2780558290","wikidata":"https://www.wikidata.org/wiki/Q134491","display_name":"Gasterosteus","level":3,"score":0.6926000118255615},{"id":"https://openalex.org/C2778334786","wikidata":"https://www.wikidata.org/wiki/Q1586270","display_name":"Variation (astronomy)","level":2,"score":0.5723000168800354},{"id":"https://openalex.org/C188382862","wikidata":"https://www.wikidata.org/wiki/Q170430","display_name":"Predation","level":2,"score":0.4325999915599823},{"id":"https://openalex.org/C2909208804","wikidata":"https://www.wikidata.org/wiki/Q127282","display_name":"Fish <Actinopterygii>","level":2,"score":0.39419999718666077},{"id":"https://openalex.org/C2908647359","wikidata":"https://www.wikidata.org/wiki/Q2625603","display_name":"Population","level":2,"score":0.3767000138759613},{"id":"https://openalex.org/C2779448149","wikidata":"https://www.wikidata.org/wiki/Q191797","display_name":"Aggression","level":2,"score":0.36649999022483826},{"id":"https://openalex.org/C18903297","wikidata":"https://www.wikidata.org/wiki/Q7150","display_name":"Ecology","level":1,"score":0.35179999470710754},{"id":"https://openalex.org/C117773394","wikidata":"https://www.wikidata.org/wiki/Q852569","display_name":"Fish migration","level":3,"score":0.3456999957561493},{"id":"https://openalex.org/C180872759","wikidata":"https://www.wikidata.org/wiki/Q874405","display_name":"Social group","level":2,"score":0.321399986743927},{"id":"https://openalex.org/C90805937","wikidata":"https://www.wikidata.org/wiki/Q7703794","display_name":"Territoriality","level":2,"score":0.31850001215934753},{"id":"https://openalex.org/C130064352","wikidata":"https://www.wikidata.org/wiki/Q853725","display_name":"Social relation","level":2,"score":0.3151000142097473},{"id":"https://openalex.org/C15744967","wikidata":"https://www.wikidata.org/wiki/Q9418","display_name":"Psychology","level":0,"score":0.30140000581741333},{"id":"https://openalex.org/C52820260","wikidata":"https://www.wikidata.org/wiki/Q7551312","display_name":"Social preferences","level":2,"score":0.2953000068664551},{"id":"https://openalex.org/C77805123","wikidata":"https://www.wikidata.org/wiki/Q161272","display_name":"Social psychology","level":1,"score":0.2922999858856201},{"id":"https://openalex.org/C149923435","wikidata":"https://www.wikidata.org/wiki/Q37732","display_name":"Demography","level":1,"score":0.2892000079154968},{"id":"https://openalex.org/C86803240","wikidata":"https://www.wikidata.org/wiki/Q420","display_name":"Biology","level":0,"score":0.2815999984741211},{"id":"https://openalex.org/C25032326","wikidata":"https://www.wikidata.org/wiki/Q185467","display_name":"Socialization","level":2,"score":0.27720001339912415},{"id":"https://openalex.org/C79416737","wikidata":"https://www.wikidata.org/wiki/Q2305519","display_name":"Social learning","level":2,"score":0.27070000767707825},{"id":"https://openalex.org/C2983406839","wikidata":"https://www.wikidata.org/wiki/Q921513","display_name":"Social behaviour","level":2,"score":0.26919999718666077},{"id":"https://openalex.org/C205649164","wikidata":"https://www.wikidata.org/wiki/Q1071","display_name":"Geography","level":0,"score":0.26330000162124634},{"id":"https://openalex.org/C2992994097","wikidata":"https://www.wikidata.org/wiki/Q7150","display_name":"Animal ecology","level":2,"score":0.25839999318122864},{"id":"https://openalex.org/C138496976","wikidata":"https://www.wikidata.org/wiki/Q175002","display_name":"Developmental psychology","level":1,"score":0.2572000026702881},{"id":"https://openalex.org/C2986275291","wikidata":"https://www.wikidata.org/wiki/Q839578","display_name":"Group behavior","level":2,"score":0.2565000057220459}],"mesh":[],"locations_count":1,"locations":[{"id":"doi:10.64898/2026.09.17.752332","doi":"https://doi.org/10.64898/2026.09.17.752332","is_oa":true,"landing_page_url":"https://doi.org/10.64898/2026.09.17.752332","pdf_url":"https://www.biorxiv.org/content/biorxiv/early/2026/09/23/2026.09.17.752332.full.pdf","source":{"id":"https://openalex.org/S4306402567","display_name":"bioRxiv (Cold Spring Harbor Laboratory)","issn_l":null,"issn":null,"is_oa":false,"is_in_doaj":false,"is_core":false,"listed_in":[],"host_organization":"https://openalex.org/I2750212522","host_organization_name":"Cold Spring Harbor Laboratory","host_organization_lineage":["https://openalex.org/I2750212522"],"host_organization_lineage_names":[],"type":"repository"},"license":"cc-by-nc","license_id":"https://openalex.org/licenses/cc-by-nc","version":"acceptedVersion","is_accepted":true,"is_published":false,"raw_source_name":"bioRxiv","raw_type":"posted-content"}],"best_oa_location":{"id":"doi:10.64898/2026.09.17.752332","doi":"https://doi.org/10.64898/2026.09.17.752332","is_oa":true,"landing_page_url":"https://doi.org/10.64898/2026.09.17.752332","pdf_url":"https://www.biorxiv.org/content/biorxiv/early/2026/09/23/2026.09.17.752332.full.pdf","source":{"id":"https://openalex.org/S4306402567","display_name":"bioRxiv (Cold Spring Harbor Laboratory)","issn_l":null,"issn":null,"is_oa":false,"is_in_doaj":false,"is_core":false,"listed_in":[],"host_organization":"https://openalex.org/I2750212522","host_organization_name":"Cold Spring Harbor Laboratory","host_organization_lineage":["https://openalex.org/I2750212522"],"host_organization_lineage_names":[],"type":"repository"},"license":"cc-by-nc","license_id":"https://openalex.org/licenses/cc-by-nc","version":"acceptedVersion","is_accepted":true,"is_published":false,"raw_source_name":"bioRxiv","raw_type":"posted-content"},"sustainable_development_goals":[],"study_designs":[],"awards":[{"id":"https://openalex.org/G140201441","display_name":"Post-Pandemic Economic Outlook: Automation, Reshoring, Education, and Growth","funder_award_id":"200914","funder_id":"https://openalex.org/F4320320924","funder_display_name":"Schweizerischer Nationalfonds zur F\u00f6rderung der Wissenschaftlichen Forschung"},{"id":"https://openalex.org/G3408928432","display_name":"Mechanisms underlying division of labour in a vertebrate model","funder_award_id":"207448","funder_id":"https://openalex.org/F4320320924","funder_display_name":"Schweizerischer Nationalfonds zur F\u00f6rderung der Wissenschaftlichen Forschung"},{"id":"https://openalex.org/G5468311820","display_name":"The eco-evo-devo of social personalities","funder_award_id":"ALWOP.668","funder_id":"https://openalex.org/F4320321800","funder_display_name":"Nederlandse Organisatie voor Wetenschappelijk Onderzoek"},{"id":"https://openalex.org/G6473417075","display_name":null,"funder_award_id":"2020GR040","funder_id":"https://openalex.org/F4320324429","funder_display_name":"Gratama Stichting"},{"id":"https://openalex.org/G8226984716","display_name":null,"funder_award_id":"01755849/WF-2019/200914","funder_id":"https://openalex.org/F4320324066","funder_display_name":"Waddenfonds"}],"funders":[{"id":"https://openalex.org/F4320320924","display_name":"Schweizerischer Nationalfonds zur F\u00f6rderung der Wissenschaftlichen Forschung","ror":"https://ror.org/00yjd3n13"},{"id":"https://openalex.org/F4320320933","display_name":"Rijksuniversiteit Groningen","ror":"https://ror.org/012p63287"},{"id":"https://openalex.org/F4320321800","display_name":"Nederlandse Organisatie voor Wetenschappelijk Onderzoek","ror":"https://ror.org/04jsz6e67"},{"id":"https://openalex.org/F4320324066","display_name":"Waddenfonds","ror":"https://ror.org/02h1t6f69"},{"id":"https://openalex.org/F4320324429","display_name":"Gratama Stichting","ror":"https://ror.org/0151rzq22"}],"has_content":{"grobid_xml":true,"pdf":true},"content_urls":{"pdf":"https://content.openalex.org/works/W7214099504.pdf","grobid_xml":"https://content.openalex.org/works/W7214099504.grobid-xml"},"referenced_works_count":0,"referenced_works":[],"related_works":["https://openalex.org/W2124730043","https://openalex.org/W4401477874","https://openalex.org/W2539498184","https://openalex.org/W2611714591","https://openalex.org/W2155563299","https://openalex.org/W4293246587","https://openalex.org/W2095065759","https://openalex.org/W4362602123","https://openalex.org/W1995823473","https://openalex.org/W2788597159"],"abstract_inverted_index":{"Abstract":[0],"Schooling":[1],"is":[2,19,106,240],"a":[3,301],"widespread":[4],"behaviour":[5,81,105,186,264,283,324,356,374],"among":[6,38,69],"pelagic":[7],"fish":[8,314],"that":[9,205,229,256,311,363,376],"reduces":[10],"locomotion":[11,272],"costs":[12,270],"of":[13,27,40,46,82,217,271,279,367,384],"migration":[14,320,386],"and":[15,25,37,68,85,90,110,143,174,192,222,273,281,291,339,375],"predation":[16,274],"risk.":[17,275],"It":[18],"characterised":[20],"by":[21,287,296,345],"coordinated":[22],"movements,":[23],"speed":[24],"orientation":[26],"group":[28,189,259],"members.":[29,260],"Although":[30],"schooling":[31,56,80,104,120,163,185,194,263,280,282,323,336,341,355],"can":[32,208,257],"vary":[33,65],"both":[34],"between":[35,66],"populations":[36,67,89,159,228,315],"individuals":[39,70,132,180],"the":[41,79,157,188,215,277,288,382],"same":[42],"population,":[43],"these":[44],"levels":[45,239],"variation":[47,97,235,347],"are":[48,112,285,293,369],"seldom":[49],"considered":[50],"together.":[51],"Here,":[52],"we":[53],"investigated":[54],"whether":[55,92,103,308],"tendencies":[57],"in":[58,98,113,154,167,198,220,227,303,348,387],"three-spined":[59,304],"sticklebacks":[60,305],"(":[61],"Gasterosteus":[62],"aculeatus":[63],")":[64],"within-populations":[71],"depending":[72],"on":[73],"their":[74,184,318,327,340,354],"phenotype.":[75],"We":[76,100,299],"first":[77],"compared":[78,325],"anadromous":[83,329],"(\u2018migrants\u2019)":[84],"recently":[86],"landlocked":[87],"(\u2018residents\u2019)":[88,321],"tested":[91],"individual":[93,218,346],"sociability":[94,219],"explained":[95],"within-population":[96],"schooling.":[99],"then":[101],"studied":[102],"modulated":[107],"when":[108,200,357],"migrants":[109,199],"residents":[111,123],"mixed":[114,358],"groups.":[115,178],"Migrant":[116],"groups":[117],"displayed":[118,133],"better":[119,137,334],"performance":[121,164],"than":[122,146,337],"with":[124,169,268,359],"higher":[125],"alignment.":[126],"Within":[127],"migrant":[128],"groups,":[129],"less":[130,145],"social":[131,148,190,246,254],"more":[134,147],"leadership,":[135],"were":[136,152,160,196,332,343],"aligned,":[138],"swam":[139],"closer":[140],"to":[141,187,243,266,306,316,326,371],"others":[142],"fatigued":[144],"fish.":[149],"These":[150,202],"patterns":[151,195,342],"absent":[153],"residents.":[155],"When":[156],"two":[158],"mixed,":[161],"overall":[162],"declined,":[165],"particularly":[166],"migrants,":[168],"schools":[170],"exhibiting":[171],"reduced":[172,269],"compaction":[173],"alignment,":[175],"resembling":[176],"resident":[177],"Thus,":[179],"did":[181,351],"not":[182,352],"modulate":[183],"composition":[191],"phenotype-dependent":[193],"disrupted":[197],"mixed.":[201],"findings":[203],"suggest":[204],"migration-related":[206,373],"behaviours":[207,255],"disappear":[209],"quickly":[210],"post-isolation.":[211],"They":[212],"also":[213],"highlight":[214],"importance":[216],"self-organisation":[221],"shaping":[223],"collective":[224],"behaviour,":[225],"especially":[226],"retain":[230],"migratory":[231],"behaviour.":[232],"Considering":[233],"behavioural":[234],"across":[236],"multiple":[237],"hierarchical":[238],"thus":[241,292],"key":[242],"understanding":[244],"complex":[245,253],"systems.":[247],"Significance":[248],"statement":[249],"Many":[250],"animals":[251],"exhibit":[252],"benefit":[258],"In":[261],"fish,":[262],"enables":[265],"migrate":[267],"Yet,":[276],"benefits":[278],"itself":[284],"shaped":[286,344],"environmental":[289],"conditions,":[290],"strongly":[294],"influenced":[295],"human":[297],"disturbance.":[298],"conducted":[300],"study":[302],"test":[307],"man-made":[309],"barriers":[310],"prohibit":[312],"some":[313],"undergo":[317],"seasonal":[319],"altered":[322],"\u2018migrant\u2019":[328],"ancestors.":[330],"Migrants":[331],"indeed":[333],"at":[335],"residents,":[338],"sociability.":[349],"Residents":[350],"improve":[353],"migrants.":[360],"This":[361],"suggests":[362],"\u223c":[364],"50":[365],"years":[366],"isolation":[368],"sufficient":[370],"losing":[372],"river":[377],"reconnection":[378],"will":[379],"likely":[380],"promote":[381],"emergence":[383],"partial":[385],"this":[388],"system.":[389]},"counts_by_year":[],"updated_date":"2026-10-28T10:33:24.094632","created_date":"2026-09-24T00:00:00"}