title: Toward major evolutionary transitions theory 2.0.
abstract: The impressive body of work on the major evolutionary transitions in the last 20 y calls for a reconstruction of the theory although a 2D account (evolution of informational systems and transitions in individuality) remains. Significant advances include the concept of fraternal and egalitarian transitions (lower-level units like and unlike, respectively). Multilevel selection, first without, then with, the collectives in focus is an important explanatory mechanism. Transitions are decomposed into phases of origin, maintenance, and transformation (i.e., further evolution) of the higher level units, which helps reduce the number of transitions in the revised list by two so that it is less top-heavy. After the transition, units show strong cooperation and very limited realized conflict. The origins of cells, the emergence of the genetic code and translation, the evolution of the eukaryotic cell, multicellularity, and the origin of human groups with language are reconsidered in some detail in the light of new data and considerations. Arguments are given why sex is not in the revised list as a separate transition. Some of the transitions can be recursive (e.g., plastids, multicellularity) or limited (transitions that share the usual features of major transitions without a massive phylogenetic impact, such as the micro- and macronuclei in ciliates). During transitions, new units of reproduction emerge, and establishment of such units requires high fidelity of reproduction (as opposed to mere replication).
authors.affiliation: Center for the Conceptual Foundations of Science, Parmenides Foundation, D-82049 Munich, Germany; Department of Plant Systematics, Ecology and Theoretical Biology, Biological Institute, Eötvös University, H-1117 Budapest, Hungary; and MTA-ELTE Theoretical Biology and Evolutionary Ecology Research Group, H-1117 Budapest, Hungary szathmary.eors@gmail.com.
authors.forename: Eörs
authors.initials: E
authors.lastname: Szathmáry
mesh_terms: D000818:Animals; D001419:Bacteria / Q000235:genetics; D005075:Biological Evolution; D003466:Cultural Characteristics; D017753:Ecosystem; D005057:Eukaryotic Cells / Q000166:cytology; D005260:Female; D022761:Gene Transfer, Horizontal; D005815:Genetic Code; D006801:Humans; D007802:Language; D008297:Male; D008928:Mitochondria / Q000235:genetics; D001678:Organelle Biogenesis; D010587:Phagocytosis; D010802:Phylogeny; D018087:Plastids / Q000235:genetics; D015203:Reproducibility of Results; D013269:Stochastic Processes
keywords: aggregative unit formation; egalitarian transitions; fraternal transitions; multilevel selection; recursive transitions
references.citation: Philos Trans R Soc Lond B Biol Sci. 2009 Nov 12;364(1533):3143-55
references.pmid: 19805423
references.citation: J Mol Evol. 2009 Nov;69(5):406-29
references.pmid: 19795157
references.citation: G3 (Bethesda). 2014 Feb 19;4(2):325-39
references.pmid: 24347631
references.citation: Proc Natl Acad Sci U S A. 1983 May;80(10):2985-9
references.pmid: 16593312
references.citation: Cold Spring Harb Perspect Biol. 2014 Apr 01;6(4):a016188
references.pmid: 24691961
references.citation: Trends Ecol Evol. 2005 Jul;20(7):363-4
references.pmid: 16701397
references.citation: Curr Opin Microbiol. 2014 Dec;22:38-48
references.pmid: 25306530
references.citation: Nature. 1992 Jul 16;358(6383):203-9
references.pmid: 1630488
references.citation: Dev Genes Evol. 2006 Dec;216(12):743-54
references.pmid: 16983540
references.citation: J Theor Biol. 2000 Sep 7;206(1):81-9
references.pmid: 10968939
references.citation: J Theor Biol. 2012 Apr 21;299:9-20
references.pmid: 21723299
references.citation: J Theor Biol. 2006 Mar 21;239(2):236-46
references.pmid: 16236324
references.citation: Proc Natl Acad Sci U S A. 1975 Jan;72(1):143-6
references.pmid: 1054490
references.citation: Orig Life Evol Biosph. 1991;21(3):165-76
references.pmid: 1724560
references.citation: Curr Biol. 2012 Aug 7;22(15):R601-2
references.pmid: 22877780
references.citation: Trends Genet. 1991 May;7(5):145-8
references.pmid: 2068786
references.citation: J Theor Biol. 1992 Nov 7;159(1):99-109
references.pmid: 1291813
references.citation: Proc Natl Acad Sci U S A. 1993 Nov 1;90(21):9916-20
references.pmid: 8234335
references.citation: J Theor Biol. 2013 Sep 7;332:171-80
references.pmid: 23648187
references.citation: J Theor Biol. 1994 Oct 7;170(3):301-9
references.pmid: 7996858
references.citation: J Mol Evol. 1999 Jun;48(6):779-83
references.pmid: 10229582
references.citation: Genetics. 2006 Dec;174(4):2173-80
references.pmid: 17028325
references.citation: BMC Evol Biol. 2011 Sep 20;11:261
references.pmid: 21933413
references.citation: Nature. 2012 Nov 1;491(7422):72-7
references.pmid: 23075853
references.citation: J Theor Biol. 1993 Oct 21;164(4):437-46
references.pmid: 8264246
references.citation: J Theor Biol. 2014 May 21;349:22-31
references.pmid: 24486248
references.citation: Biol Direct. 2012 Oct 31;7:38; discussion 38
references.pmid: 23114029
references.citation: Acc Chem Res. 2012 Dec 18;45(12):2097-105
references.pmid: 22891822
references.citation: Nature. 2010 Oct 21;467(7318):929-34
references.pmid: 20962839
references.citation: Orig Life Evol Biosph. 2003 Oct;33(4-5):405-32
references.pmid: 14604184
references.citation: Ann N Y Acad Sci. 2015 Apr;1341:75-95
references.pmid: 25735569
references.citation: Angew Chem Int Ed Engl. 2003 Jun 30;42(25):2909-12
references.pmid: 12833356
references.citation: J Theor Biol. 2015 Apr 21;371:45-58
references.pmid: 25681798
references.citation: Proc Biol Sci. 1999 Aug 7;266(1428):1571-7
references.pmid: 10467746
references.citation: Int J Biochem Cell Biol. 2009 Feb;41(2):307-22
references.pmid: 18935970
references.citation: Philos Trans R Soc Lond B Biol Sci. 2011 Oct 27;366(1580):2902-9
references.pmid: 21930581
references.citation: Nature. 1979 Aug 9;280(5722):445-6
references.pmid: 460422
references.citation: Biol Direct. 2011 Jun 30;6:35
references.pmid: 21714941
references.citation: Q Rev Biol. 2007 Dec;82(4):327-48
references.pmid: 18217526
references.citation: Res Microbiol. 2009 Sep;160(7):473-80
references.pmid: 19647074
references.citation: BMC Biol. 2010 Mar 10;8:21
references.pmid: 20219099
references.citation: Trends Ecol Evol. 1989 Jul;4(7):200-4
references.pmid: 21227350
references.citation: Trends Ecol Evol. 1991 Dec;6(12):407-8
references.pmid: 21232522
references.citation: J Mol Evol. 1995 May;40(5):545-50
references.pmid: 7783229
references.citation: Nat Rev Genet. 2005 Sep;6(9):699-708
references.pmid: 16151375
references.citation: Nature. 2011 Jan 20;469(7330):393-6
references.pmid: 21248849
references.citation: J Hum Evol. 1999 May;36(5):461-85
references.pmid: 10222165
references.citation: Biol Direct. 2007 Jan 19;2:3
references.pmid: 17239231
references.citation: Biol Direct. 2012 Jan 05;7:1; discussion 1
references.pmid: 22221860
references.citation: Cold Spring Harb Perspect Biol. 2014 Jul 18;6(12):a015990
references.pmid: 25038049
references.citation: Int J Mol Sci. 2011;12(5):3085-101
references.pmid: 21686171
references.citation: Proc Natl Acad Sci U S A. 1989 Sep;86(18):7054-8
references.pmid: 2476811
references.citation: Integr Comp Biol. 2003 Feb;43(1):64-73
references.pmid: 21680410
references.citation: Trends Ecol Evol. 2005 Dec;20(12):650
references.pmid: 16701451
references.citation: Nature. 1995 Mar 16;374(6519):227-32
references.pmid: 7885442
references.citation: J Theor Biol. 1987 Oct 21;128(4):463-86
references.pmid: 2451771
references.citation: Proc Natl Acad Sci U S A. 2002 Jun 25;99(13):8742-7
references.pmid: 12077305
references.citation: J Theor Biol. 1986 Mar 7;119(1):1-24
references.pmid: 3713221
references.citation: Proc Natl Acad Sci U S A. 2006 Jul 11;103(28):10696-701
references.pmid: 16818880
references.citation: Naturwissenschaften. 1971 Oct;58(10):465-523
references.pmid: 4942363
references.citation: Nature. 2002 Nov 21;420(6913):340-3
references.pmid: 12447445
references.citation: Trends Ecol Evol. 1991 Aug;6(8):257-62
references.pmid: 21232473
references.citation: J Mol Evol. 2009 Nov;69(5):458-69
references.pmid: 19806387
references.citation: Integr Comp Biol. 2003 Feb;43(1):82-6
references.pmid: 21680412
references.citation: Proc Natl Acad Sci U S A. 2012 Apr 3;109(14):5340-5
references.pmid: 22371600
references.citation: Proc Natl Acad Sci U S A. 1992 Apr 1;89(7):2614-8
references.pmid: 1372984
references.citation: Front Microbiol. 2014 Jun 13;5:274
references.pmid: 24982654