a92b8b7881e3b77c0494973cf0192277 12915_2023_Article_1716.pdf f81705bec8dd239cac0bdaafe56ff4b0e2bd26ae 12915_2023_Article_1716.pdf 739112149ea37f04c19f92dcbfb69a7a1bd0b0a4992bd4e9f5d07af4e1e630f8 12915_2023_Article_1716.pdf Title: Directional selection coupled with kin selection favors the establishment of senescence Subject: BMC Biology, https://doi.org/10.1186/s12915-023-01716-w Keywords: Aging;Evolution;Longevity;Evolvability;Kin selection Author: András Szilágyi Creator: Adobe InDesign 15.1 (Windows) Producer: Adobe PDF Library 15.0; modified using iText® 5.3.5 ©2000-2012 1T3XT BVBA (SPRINGER SBM; licensed version) CreationDate: Sat Oct 7 11:42:07 2023 CEST ModDate: Sun Nov 19 03:53:39 2023 CET Custom Metadata: yes Metadata Stream: yes Tagged: no UserProperties: no Suspects: no Form: none JavaScript: no Pages: 14 Encrypted: no Page size: 595.276 x 790.866 pts Page rot: 0 File size: 3682536 bytes Optimized: no PDF version: 1.4 name type encoding emb sub uni object ID 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12915_2023_Article_1716.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-06-29 15:35:02 CEST Size: 3682536 Format: PDF Version: 1.4 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf PDFMetadata: Objects: 1447 FreeObjects: 1 IncrementalUpdates: 0 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: false NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseOutlines Language: en-US Outlines: Item: Title: Directional selection coupled with kin selection favors the establishment of senescence Children: Item: Title: Abstract Children: Item: Title: Background Item: Title: Results Item: Title: Conclusions Item: Title: Background Item: Title: Results Children: Item: Title: Spatial dynamics Item: Title: Lifespan and fecundity evolution under directional and stabilizing selection Item: Title: Kin selection Item: Title: Parameter screening Children: Item: Title: Mutation Item: Title: Recombination Item: Title: Invasibility analysis Item: Title: The effect of baseline (extrinsic) mortality Item: Title: Aging under directional selection is due to its indirect effect on evolvability Item: Title: The effect of spatial mixing Item: Title: Discussion Item: Title: Conclusions Item: Title: Methods Children: Item: Title: Death rate Item: Title: Fecundity Item: Title: Dynamics Item: Title: Spatial mixing Item: Title: Anchor 25 Item: Title: Acknowledgements Item: Title: References Info: Title: Directional selection coupled with kin selection favors the establishment of senescence Author: András Szilágyi Subject: BMC Biology, https://doi.org/10.1186/s12915-023-01716-w Keywords: Aging;Evolution;Longevity;Evolvability;Kin selection Creator: Adobe InDesign 15.1 (Windows) Producer: Adobe PDF Library 15.0; modified using iText® 5.3.5 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JWLHXL+MTSYN19 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+MTSYN20 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN20 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-It24 FontSubset: true FirstChar: 27 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It24 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI34 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI34 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI33 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI33 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: MyriadPro-Light FirstChar: 0 LastChar: 255 FontDescriptor: FontName: MyriadPro-Light Flags: Nonsymbolic FontBBox: -49, -250, 1046, 883 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-It23 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It23 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It4 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It4 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular22 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular22 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: WarnockPro-It3 FirstChar: 0 LastChar: 255 FontDescriptor: FontName: WarnockPro-It3 Flags: Serif, Nonsymbolic, Italic FontBBox: -221, -533, 1474, 950 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-Regular2 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular2 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It2 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It2 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular3 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular3 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI2 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI2 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt9 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt9 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-Light6 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-Light6 Flags: Symbolic, SmallCap FontBBox: 0, -11, 500, 721 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN5 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN5 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: EYOKVV+MyriadPro-LightIt11 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt11 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI10 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI10 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt10 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt10 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MTSYN4 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: EYOKVV+MTSYN4 Flags: Symbolic, AllCap FontBBox: 0, -156, 299, 662 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI11 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI11 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI8 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI8 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-Light8 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-Light8 Flags: Symbolic, SmallCap FontBBox: 0, -11, 500, 721 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI4 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI4 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt13 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt13 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-Light7 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-Light7 Flags: Symbolic, SmallCap FontBBox: 0, -11, 500, 721 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI9 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI9 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt12 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt12 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular4 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular4 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI7 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI7 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MTSYN3 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: EYOKVV+MTSYN3 Flags: Symbolic, AllCap FontBBox: 0, -156, 299, 662 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: DRHXOS+MyriadPro-LightIt5 FontSubset: true FirstChar: 0 LastChar: 255 FontDescriptor: FontName: DRHXOS+MyriadPro-LightIt5 Flags: Nonsymbolic, Italic FontBBox: -121, -250, 1007, 863 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: EYOKVV+MyriadPro-LightIt4 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt4 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI5 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI5 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-Light5 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-Light5 Flags: Symbolic, SmallCap FontBBox: 0, -11, 500, 721 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt8 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt8 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt7 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt7 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt6 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt6 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI6 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI6 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI3 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI3 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-Light4 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-Light4 Flags: Symbolic, SmallCap FontBBox: 0, -11, 500, 721 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt3 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt3 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MTSYN FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MTSYN Flags: Symbolic, AllCap FontBBox: 0, -156, 299, 662 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: MyriadPro-Semibold FirstChar: 0 LastChar: 255 FontDescriptor: FontName: MyriadPro-Semibold Flags: Nonsymbolic FontBBox: -161, -250, 1198, 972 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: EYOKVV+MyriadPro-Light3 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-Light3 Flags: Symbolic, SmallCap FontBBox: 0, -11, 500, 721 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-LightIt2 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-LightIt2 Flags: Symbolic FontBBox: -9, -198, 616, 710 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MTSYN2 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: EYOKVV+MTSYN2 Flags: Symbolic, AllCap FontBBox: 0, -156, 299, 662 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: EYOKVV+MyriadPro-Light2 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: EYOKVV+MyriadPro-Light2 Flags: Symbolic, SmallCap FontBBox: 0, -11, 500, 721 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular26 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular26 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN23 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN23 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: HCQFHP+CMEX103 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: HCQFHP+CMEX103 Flags: Symbolic FontBBox: 0, -2359, 999, 722 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It27 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It27 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI36 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI36 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN25 FontSubset: true FirstChar: 28 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN25 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-It25 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It25 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It30 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It30 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI41 FontSubset: true FirstChar: 28 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI41 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: HCQFHP+CMEX104 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: HCQFHP+CMEX104 Flags: Symbolic FontBBox: 0, -2359, 999, 722 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular25 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular25 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It26 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It26 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular24 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular24 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular29 FontSubset: true FirstChar: 24 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular29 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN22 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN22 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: HCQFHP+CMEX105 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: HCQFHP+CMEX105 Flags: Symbolic FontBBox: 0, -2359, 999, 722 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular28 FontSubset: true FirstChar: 27 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular28 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN24 FontSubset: true FirstChar: 27 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN24 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-It29 FontSubset: true FirstChar: 28 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It29 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI38 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI38 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN53 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN53 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-It32 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It32 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It68 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It68 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN26 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN26 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-Regular59 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular59 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI37 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI37 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular30 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular30 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI40 FontSubset: true FirstChar: 29 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI40 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: HCQFHP+STIX-Regular FontSubset: true FirstChar: 31 LastChar: 216 FontDescriptor: FontName: HCQFHP+STIX-Regular Flags: Serif, Nonsymbolic FontBBox: -970, -486, 2208, 1023 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: 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BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN7 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN7 Flags: Symbolic FontBBox: 0, -181, 724, 680 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: JWLHXL+WarnockPro-It5 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It5 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular6 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular6 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTSYN6 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTSYN6 Flags: Symbolic FontBBox: 0, -181, 724, 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FontDescriptor: FontName: JWLHXL+MTMI48 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+MTMI18 FontSubset: true FirstChar: 30 LastChar: 32 FontDescriptor: FontName: JWLHXL+MTMI18 Flags: Symbolic FontBBox: 0, -207, 683, 676 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular36 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular36 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It9 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It9 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It37 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It37 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It7 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It7 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-It36 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-It36 Flags: Symbolic FontBBox: -191, -240, 821, 709 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true Font: BaseFont: JWLHXL+WarnockPro-Regular8 FontSubset: true FirstChar: 31 LastChar: 32 FontDescriptor: FontName: JWLHXL+WarnockPro-Regular8 Flags: Symbolic, SmallCap FontBBox: 0, -169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: 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-169, 824, 712 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true XMP: 2023-10-07T09:42:07 Adobe InDesign 15.1 (Windows) 2023-11-19T03:53:39+01:00 2023-11-19T03:53:39+01:00 Adobe PDF Library 15.0; modified using iText® 5.3.5 ©2000-2012 1T3XT BVBA (SPRINGER SBM; licensed version) Aging;Evolution;Longevity;Evolvability;Kin selection False application/pdf https://doi.org/10.1186/s12915-023-01716-w BioMed Central BMC Biology, https://doi.org/10.1186/s12915-023-01716-w Aging Evolution Longevity Evolvability Kin selection Directional selection coupled with kin selection favors the establishment of senescence András Szilágyi Tamás Czárán Mauro Santos Eörs Szathmáry 10.1186/s12915-023-01716-w 2010-04-23 true springer.com springerlink.com https://doi.org/10.1186/s12915-023-01716-w 10.1186/s12915-023-01716-w 1741-7007 journal BMC Biology The Author(s) 2010-04-23 true 10.1186/s12915-023-01716-w noindex springer.com springerlink.com uuid:8f4e7c34-e7a8-490a-a75d-2fd7fcbc5768 uuid:144b8dd3-72f1-4950-9981-30984a62435f default 1 converted uuid:8f4e7c34-e7a8-490a-a75d-2fd7fcbc5768 converted to PDF/A-2b pdfToolbox 2023-10-07T12:23:21Z 2 B Eörs Szathmáry http://orcid.org/0000-0001-5227-2997 http://ns.adobe.com/pdfx/1.3/ pdfx Adobe Document Info PDF eXtension Schema external Mirrors crossmark:MajorVersionDate CrossmarkMajorVersionDate Text external Mirrors crossmark:CrossmarkDomainExclusive CrossmarkDomainExclusive Text internal Mirrors crossmark:DOI doi Text external Mirrors crossmark:CrosMarkDomains CrossMarkDomains seq Text internal A name object indicating whether the document has been modified to include trapping information robots Text internal ID of PDF/X standard GTS_PDFXVersion Text internal Conformance level of PDF/X standard GTS_PDFXConformance Text internal Company creating the PDF Company Text internal Date when document was last modified SourceModified Text http://crossref.org/crossmark/1.0/ crossmark Crossmark Schema internal Usual same as prism:doi DOI Text external The date when a publication was publishe. MajorVersionDate Text internal CrossmarkDomainExclusive CrossmarkDomainExclusive Text internal CrossMarkDomains CrossMarkDomains seq Text http://prismstandard.org/namespaces/basic/2.0/ prism Prism Schema external This element provides the url for an article or unit of content. The attribute platform is optionally allowed for situations in which multiple URLs must be specified. PRISM recommends that a subset of the PCV platform values, namely “mobile” and “web”, be used in conjunction with this element. NOTE: PRISM recommends against the use of the #other value allowed in the PRISM Platform controlled vocabulary. In lieu of using #other please reach out to the PRISM group at prism-wg@yahoogroups.com to request addition of your term to the Platform Controlled Vocabulary. url URI external The Digital Object Identifier for the article. The DOI may also be used as the dc:identifier. If used as a dc:identifier, the URI form should be captured, and the bare identifier should also be captured using prism:doi. If an alternate unique identifier is used as the required dc:identifier, then the DOI should be specified as a bare identifier within prism:doi only. If the URL associated with a DOI is to be specified, then prism:url may be used in conjunction with prism:doi in order to provide the service endpoint (i.e. the URL). doi Text external ISSN for an electronic version of the issue in which the resource occurs. Permits publishers to include a second ISSN, identifying an electronic version of the issue in which the resource occurs (therefore e(lectronic)Issn. If used, prism:eIssn MUST contain the ISSN of the electronic version. issn Text internal Volume number volume Text internal Issue number number Text internal Starting page startingPage Text internal Ending page endingPage Text external The aggregation type specifies the unit of aggregation for a content collection. Comment PRISM recommends that the PRISM Aggregation Type Controlled Vocabulary be used to provide values for this element. Note: PRISM recommends against the use of the #other value currently allowed in this controlled vocabulary. In lieu of using #other please reach out to the PRISM group at info@prismstandard.org to request addition of your term to the Aggregation Type Controlled Vocabulary. aggregationType Text external Title of the magazine, or other publication, in which a resource was/will be published. Typically this will be used to provide the name of the magazine an article appeared in as metadata for the article, along with information such as the article title, the publisher, volume, number, and cover date. Note: Publication name can be used to differentiate between a print magazine and the online version if the names are different such as “magazine” and “magazine.com.” publicationName Text external Copyright copyright Text http://ns.adobe.com/pdf/1.3/ pdf Adobe PDF Schema internal A name object indicating whether the document has been modified to include trapping information Trapped Text http://ns.adobe.com/xap/1.0/mm/ xmpMM XMP Media Management Schema internal UUID based identifier for specific incarnation of a document InstanceID URI internal The common identifier for all versions and renditions of a document. DocumentID URI internal The common identifier for all versions and renditions of a document. OriginalDocumentID URI http://www.aiim.org/pdfa/ns/id/ pdfaid PDF/A ID Schema internal Part of PDF/A standard part Integer internal Amendment of PDF/A standard amd Text internal Conformance level of PDF/A standard conformance Text Springer Nature ORCID Schema http://springernature.com/ns/xmpExtensions/2.0/ sn authorInfo Bag AuthorInformation external Author information: contains the name of each author and his/her ORCiD (ORCiD: Open Researcher and Contributor ID). An ORCiD is a persistent identifier (a non-proprietary alphanumeric code) to uniquely identify scientific and other academic authors. AuthorInformation http://springernature.com/ns/xmpExtensions/2.0/authorInfo/ author Specifies the types of author information: name and ORCID of an author. name Text Gives the name of an author. orcid URI Gives the ORCID of an author. 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