{"id":"https://openalex.org/W2317421091","doi":"https://doi.org/10.7551/978-0-262-33027-5-ch091","title":"Simpson\u2019s Paradox, Co-operation and Individuality in Bacterial Biofilms","display_name":"Simpson\u2019s Paradox, Co-operation and Individuality in Bacterial Biofilms","publication_year":2015,"publication_date":"2015-07-18","ids":{"openalex":"https://openalex.org/W2317421091","doi":"https://doi.org/10.7551/978-0-262-33027-5-ch091","mag":"2317421091"},"language":"en","primary_location":{"id":"doi:10.7551/978-0-262-33027-5-ch091","is_oa":true,"landing_page_url":"https://doi.org/10.7551/978-0-262-33027-5-ch091","pdf_url":"http://www.mitpressjournals.org/doi/pdf/10.1162/978-0-262-33027-5-ch091","source":null,"license":null,"license_id":null,"version":"publishedVersion","is_accepted":true,"is_published":true,"raw_source_name":"07/20/2015-07/24/2015","raw_type":"proceedings-article"},"type":"article","indexed_in":["crossref"],"open_access":{"is_oa":true,"oa_status":"gold","oa_url":"http://www.mitpressjournals.org/doi/pdf/10.1162/978-0-262-33027-5-ch091","any_repository_has_fulltext":null},"authorships":[{"author_position":"first","author":{"id":"https://openalex.org/A5046702525","display_name":"Alexandra S. Penn","orcid":"https://orcid.org/0000-0001-5107-2862"},"institutions":[{"id":"https://openalex.org/I28290843","display_name":"University of Surrey","ror":"https://ror.org/00ks66431","country_code":"GB","type":"education","lineage":["https://openalex.org/I28290843"]}],"countries":["GB"],"is_corresponding":true,"raw_author_name":"Alexandra Penn","raw_affiliation_strings":["University of Surrey"],"raw_orcid":null,"affiliations":[{"raw_affiliation_string":"University of Surrey","institution_ids":["https://openalex.org/I28290843"]}]}],"institutions":[],"countries_distinct_count":1,"institutions_distinct_count":1,"corresponding_author_ids":["https://openalex.org/A5046702525"],"corresponding_institution_ids":["https://openalex.org/I28290843"],"apc_list":null,"apc_paid":null,"fwci":0.0,"has_fulltext":false,"cited_by_count":1,"citation_normalized_percentile":{"value":0.22606454,"is_in_top_1_percent":false,"is_in_top_10_percent":false},"cited_by_percentile_year":{"min":90,"max":94},"biblio":{"volume":null,"issue":null,"first_page":"520","last_page":"520"},"is_retracted":false,"is_paratext":false,"is_xpac":false,"primary_topic":{"id":"https://openalex.org/T11252","display_name":"Evolutionary Game Theory and Cooperation","score":0.9976000189781189,"subfield":{"id":"https://openalex.org/subfields/3312","display_name":"Sociology and Political Science"},"field":{"id":"https://openalex.org/fields/33","display_name":"Social Sciences"},"domain":{"id":"https://openalex.org/domains/2","display_name":"Social Sciences"}},"topics":[{"id":"https://openalex.org/T11252","display_name":"Evolutionary Game Theory and Cooperation","score":0.9976000189781189,"subfield":{"id":"https://openalex.org/subfields/3312","display_name":"Sociology and Political Science"},"field":{"id":"https://openalex.org/fields/33","display_name":"Social Sciences"},"domain":{"id":"https://openalex.org/domains/2","display_name":"Social Sciences"}},{"id":"https://openalex.org/T11764","display_name":"Evolution and Genetic Dynamics","score":0.9764999747276306,"subfield":{"id":"https://openalex.org/subfields/1311","display_name":"Genetics"},"field":{"id":"https://openalex.org/fields/13","display_name":"Biochemistry, Genetics and Molecular Biology"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}},{"id":"https://openalex.org/T10593","display_name":"Bacterial biofilms and quorum sensing","score":0.9711999893188477,"subfield":{"id":"https://openalex.org/subfields/1312","display_name":"Molecular Biology"},"field":{"id":"https://openalex.org/fields/13","display_name":"Biochemistry, Genetics and Molecular Biology"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}}],"keywords":[{"id":"https://openalex.org/keywords/biofilm","display_name":"Biofilm","score":0.6153930425643921},{"id":"https://openalex.org/keywords/computer-science","display_name":"Computer science","score":0.4290805459022522},{"id":"https://openalex.org/keywords/environmental-science","display_name":"Environmental science","score":0.3410625755786896},{"id":"https://openalex.org/keywords/geology","display_name":"Geology","score":0.2202942967414856},{"id":"https://openalex.org/keywords/bacteria","display_name":"Bacteria","score":0.09297454357147217},{"id":"https://openalex.org/keywords/paleontology","display_name":"Paleontology","score":0.05576807260513306}],"concepts":[{"id":"https://openalex.org/C58123911","wikidata":"https://www.wikidata.org/wiki/Q467410","display_name":"Biofilm","level":3,"score":0.6153930425643921},{"id":"https://openalex.org/C41008148","wikidata":"https://www.wikidata.org/wiki/Q21198","display_name":"Computer science","level":0,"score":0.4290805459022522},{"id":"https://openalex.org/C39432304","wikidata":"https://www.wikidata.org/wiki/Q188847","display_name":"Environmental science","level":0,"score":0.3410625755786896},{"id":"https://openalex.org/C127313418","wikidata":"https://www.wikidata.org/wiki/Q1069","display_name":"Geology","level":0,"score":0.2202942967414856},{"id":"https://openalex.org/C523546767","wikidata":"https://www.wikidata.org/wiki/Q10876","display_name":"Bacteria","level":2,"score":0.09297454357147217},{"id":"https://openalex.org/C151730666","wikidata":"https://www.wikidata.org/wiki/Q7205","display_name":"Paleontology","level":1,"score":0.05576807260513306}],"mesh":[],"locations_count":1,"locations":[{"id":"doi:10.7551/978-0-262-33027-5-ch091","is_oa":true,"landing_page_url":"https://doi.org/10.7551/978-0-262-33027-5-ch091","pdf_url":"http://www.mitpressjournals.org/doi/pdf/10.1162/978-0-262-33027-5-ch091","source":null,"license":null,"license_id":null,"version":"publishedVersion","is_accepted":true,"is_published":true,"raw_source_name":"07/20/2015-07/24/2015","raw_type":"proceedings-article"}],"best_oa_location":{"id":"doi:10.7551/978-0-262-33027-5-ch091","is_oa":true,"landing_page_url":"https://doi.org/10.7551/978-0-262-33027-5-ch091","pdf_url":"http://www.mitpressjournals.org/doi/pdf/10.1162/978-0-262-33027-5-ch091","source":null,"license":null,"license_id":null,"version":"publishedVersion","is_accepted":true,"is_published":true,"raw_source_name":"07/20/2015-07/24/2015","raw_type":"proceedings-article"},"sustainable_development_goals":[{"display_name":"Partnerships for the goals","id":"https://metadata.un.org/sdg/17","score":0.6000000238418579}],"awards":[],"funders":[],"has_content":{"pdf":false,"grobid_xml":false},"content_urls":null,"referenced_works_count":14,"referenced_works":["https://openalex.org/W1604756073","https://openalex.org/W1871092878","https://openalex.org/W2056678000","https://openalex.org/W2060860079","https://openalex.org/W2066034383","https://openalex.org/W2116738092","https://openalex.org/W2122836009","https://openalex.org/W2141401164","https://openalex.org/W2146433021","https://openalex.org/W2161112276","https://openalex.org/W2163902276","https://openalex.org/W2170784505","https://openalex.org/W2753514269","https://openalex.org/W2905472553"],"related_works":["https://openalex.org/W2899084033","https://openalex.org/W2748952813","https://openalex.org/W2756188369","https://openalex.org/W2947973176","https://openalex.org/W2948279684","https://openalex.org/W2087552300","https://openalex.org/W4220932551","https://openalex.org/W4246525962","https://openalex.org/W2332685989","https://openalex.org/W2130763979"],"abstract_inverted_index":{"Bacteria":[0],"often":[1],"live":[2],"in":[3,37,119,135,143,171,191,201,214,329,415,438,440],"group":[4,130,325,342,406],"structures":[5,241],"known":[6,155],"as":[7,17,156,217,375,399,401,404],"biofilms":[8,38,398],"within":[9,128,239,388,397],"which":[10,251,344,357,452],"they":[11],"commonly":[12,36],"display":[13],"co-operative":[14],"behaviours,":[15],"such":[16],"the":[18,79,115,149,162,210,229,271,310,318,330,338,341,345,373,376,385,405,412,419,441],"production":[19,211],"of":[20,69,81,117,138,152,212,232,258,277,294,313,340,378,421,444],"public":[21,331,346],"goods":[22],"(Ghannoum":[23],"&":[24,86,224],"O\u2019Toole":[25],"2004,":[26,92,94],"Crespi":[27],"2001,":[28,43],"West":[29],"et":[30,41,45,57,90,96,206,298],"al.":[31,42,46,58,91,97,207,299],"2007).":[32,98],"Noncooperative":[33],"cheats":[34,273],"arise":[35],"(de":[39],"Vos":[40],"Schaber":[44],"2004),":[47],"but":[48,185],"despite":[49,122,140],"what":[50],"theory":[51],"might":[52],"predict,":[53],"(Hardin":[54],"1968,":[55],"Rankin":[56],"2007),":[59,226],"co-operation":[60,222,396],"does":[61],"not":[62,197],"seem":[63],"to":[64,126,417],"be":[65,359,409],"disrupted.":[66],"The":[67,256],"stability":[68],"these":[70],"behaviours":[71],"requires":[72],"explanation":[73],"and":[74,111,167,234,384,423,432,437,458],"could":[75],"cast":[76],"light":[77],"on":[78],"evolution":[80],"multi-cellularity":[82],"experimentally":[83],"(e.g.":[84],"Rainey":[85,87],"2003,":[88],"Griffin":[89],"Kreft":[93],"Buckling":[95],"Theory":[99],"tells":[100],"us":[101],"that":[102,178,315,324,337,448],"repeated":[103],"aggregation":[104],"into":[105],"local":[106,145],"groups,":[107,153,194],"interleaved":[108],"with":[109,270,280,285,372],"dispersal":[110,168],"remixing,":[112],"can":[113],"increase":[114,134],"level":[116],"cooperation":[118],"a":[120,123,141,202,218,352,362,445],"population":[121,204,382,454],"selective":[124],"disadvantage":[125],"cooperating":[127],"any":[129],"(Wilson":[131],"1980).":[132],"This":[133,355,390],"global":[136],"proportion":[137,272],"co-operators":[139],"decrease":[142],"all":[144],"proportions,":[146],"caused":[147],"by":[148,361,411],"differential":[150],"growth":[151,257,266,282],"is":[154,348,351],"Simpson\u2019s":[157,179,295,322],"paradox":[158,180,296,323],"(Simpson":[159],"1951).":[160],"Given":[161],"microcolony":[163,240,254,278],"(small":[164],"sub-group)":[165],"formation":[166],"behaviour":[169,460],"observed":[170],"natural":[172,203],"biofilms,":[173],"it":[174,187,195,335],"has":[175,188,196,391],"been":[176,189,199],"suggested":[177],"may":[181,316,407,461],"explain":[182],"bacterial":[183],"cooperation;":[184],"although":[186],"demonstrated":[190,200],"artificially":[192],"constructed":[193],"yet":[198],"(Chuang":[205],"2009).":[208],"Using":[209],"siderophores":[213],"Pseudomonas":[215],"aeruginosa":[216],"model":[219],"system":[220],"for":[221,321,367,394],"(Varma":[223],"Chincholker":[225],"we":[227,290,302],"measured":[228],"change":[230,418],"frequency":[231],"co-operator":[233],"siderophore-deficient":[235],"cheat":[236],"strains":[237],"in-situ":[238],"over":[242,253,343],"time.":[243],"We":[244],"detected":[245],"significant":[246],"within-type":[247],"negative":[248],"densitydependent":[249],"effects":[250],"vary":[252],"development.":[255],"types":[259],"was":[260,267],"self-limiting":[261],"at":[262],"different":[263],"times:":[264],"Cheat":[265],"negatively":[268,283],"correlated":[269,284],"during":[274],"early":[275],"stages":[276],"development,":[279],"wildtype":[281],"wild-type":[286],"biomass":[287],"later.":[288],"However,":[289],"found":[291],"no":[292],"evidence":[293],"(Penn":[297],"2012).":[300],"Instead":[301],"saw":[303],"clear":[304],"within-microcolony":[305,425],"spatial":[306],"structure":[307],"(cheats":[308],"occupying":[309],"interior":[311],"portions":[312],"microcolonies)":[314],"violate":[317],"assumption":[319],"required":[320],"members":[326],"share":[327],"equally":[328],"good.":[332],"In":[333],"fact,":[334],"seems":[336],"extent":[339],"good":[347],"being":[349],"shared":[350],"dynamic":[353],"entity.":[354],"group,":[356],"will":[358,428],"defined":[360],"lower":[363],"threshold":[364],"siderophore":[365],"concentration":[366],"effective":[368],"iron":[369],"chelation,":[370],"codevelops":[371],"biofilm":[374],"result":[377],"an":[379],"interaction":[380],"between":[381],"dynamics":[383],"react-diffusion":[386],"processes":[387],"it.":[389],"interesting":[392],"consequences":[393],"understanding":[395],"well":[400],"major":[402],"transitions,":[403],"potentially":[408],"influenced":[410],"bacteria":[413],"themselves":[414],"order":[416],"context":[420,443],"selection":[422],"promote":[424],"\u201cindividuality\u201d.":[426],"I":[427],"discuss":[429],"our":[430],"observations":[431],"continuing":[433],"work,":[434],"both":[435],"experimental":[436],"simulation,":[439],"broader":[442],"theoretical":[446],"framework":[447],"suggests":[449],"how":[450],"factors":[451],"affect":[453],"structure,":[455],"higher-level":[456],"individuality":[457],"cooperative":[459],"co-evolve.":[462]},"counts_by_year":[{"year":2022,"cited_by_count":1}],"updated_date":"2026-05-19T21:40:30.786675","created_date":"2016-06-24T00:00:00"}
