title: Colloquium paper: how grandmother effects plus individual variation in frailty shape fertility and mortality: guidance from human-chimpanzee comparisons.
abstract: In the first paper to present formal theory explaining that senescence is a consequence of natural selection, W. D. Hamilton concluded that human postmenopausal longevity results from the contributions of ancestral grandmothers to the reproduction of their relatives. A grandmother hypothesis, subsequently elaborated with additional lines of evidence, helps explain both exceptional longevity and additional features of life history that distinguish humans from the other great apes. However, some of the variation observed in aging rates seems inconsistent with the tradeoffs between current and future reproduction identified by theory. In humans and chimpanzees, our nearest living relatives, individuals who bear offspring at faster rates do not cease bearing sooner. They continue to be fertile longer instead. Furthermore, within both species, groups with lower overall mortality rates have faster rates of increase in death risk with advancing age. These apparent contradictions to the expected life history tradeoffs likely result from heterogeneity in frailty among individuals. Whereas robust and frail alike must allocate investments between current and future reproduction, the more robust can afford more of both. This heterogeneity, combined with evolutionary tradeoffs and the key role of ancestral grandmothers they identify, helps explain aspects of human aging that increasingly concern us all.
authors.affiliation: Department of Anthropology, University of Utah, Salt Lake City, UT 84112, USA.
authors.forename: Kristen
authors.initials: K
authors.lastname: Hawkes
mesh_terms: D000328:Adult; D000368:Aged; D000369:Aged, 80 and over; D000375:Aging; D000818:Animals; D044822:Biodiversity; D005075:Biological Evolution; D005260:Female; D006801:Humans; D008136:Longevity; D008593:Menopause; D008875:Middle Aged; D008954:Models, Biological; D008957:Models, Genetic; D008962:Models, Theoretical; D002679:Pan troglodytes; D017698:Postmenopause
references.citation: Am J Hum Biol. 2001 Jul-Aug;13(4):465-78
references.pmid: 11400217
references.citation: Am J Hum Biol. 2009 Jul-Aug;21(4):533-40
references.pmid: 19298004
references.citation: Popul Dev Rev. 2003 Sep 1;29 Supp(3):270-291
references.pmid: 19337587
references.citation: Biol Reprod. 1994 Mar;50(3):653-63
references.pmid: 8167237
references.citation: Infect Genet Evol. 2009 Dec;9(6):1371-80
references.pmid: 19804845
references.citation: Science. 1995 Oct 6;270(5233):53-9
references.pmid: 7569951
references.citation: Am J Hum Biol. 2009 Jul-Aug;21(4):578-86
references.pmid: 19213006
references.citation: Hum Nat. 2002 Jun;13(2):239-67
references.pmid: 26192759
references.citation: J Theor Biol. 1966 Sep;12(1):12-45
references.pmid: 6015424
references.citation: Proc Natl Acad Sci U S A. 2003 Aug 5;100(16):9637-42
references.pmid: 12878733
references.citation: Eugen Q. 1961 Jun;8:81-91
references.pmid: 13713432
references.citation: Proc Natl Acad Sci U S A. 2010 May 11;107 Suppl 2:8902-9
references.pmid: 20445105
references.citation: J Dev Orig Health Dis. 2010 Feb;1(1):26-34
references.pmid: 20198106
references.citation: Nature. 1997 Jul 31;388(6641):434
references.pmid: 9242401
references.citation: BMJ. 1989 Mar 4;298(6673):564-7
references.pmid: 2495113
references.citation: Am Stat. 1985 Aug;39(3):176-85
references.pmid: 12267300
references.citation: Proc Natl Acad Sci U S A. 1996 Dec 24;93(26):15249-53
references.pmid: 8986796
references.citation: Annu Rev Public Health. 2008;29:91-113
references.pmid: 18031222
references.citation: Am J Phys Anthropol. 2009 Sep;140(1):173-6
references.pmid: 19373844
references.citation: Science. 1998 May 8;280(5365):855-60
references.pmid: 9599158
references.citation: Mol Biol Evol. 2008 Jan;25(1):120-30
references.pmid: 17998254
references.citation: Philos Trans R Soc Lond B Biol Sci. 1991 Apr 29;332(1262):15-24
references.pmid: 1677205
references.citation: Am J Primatol. 2005 Oct;67(2):199-207
references.pmid: 16229006
references.citation: Nature. 1997 Sep 11;389(6647):133
references.pmid: 9296486
references.citation: J Anat. 2008 Apr;212(4):394-425
references.pmid: 18380863
references.citation: Am J Primatol. 2005 Nov;67(3):281-98
references.pmid: 16287103
references.citation: Neurobiol Aging. 2009 Apr;30(4):507-14
references.pmid: 19231028
references.citation: J Gerontol A Biol Sci Med Sci. 2002 May;57(5):B202-6
references.pmid: 11983718
references.citation: Proc Natl Acad Sci U S A. 2001 Feb 27;98(5):2934-9
references.pmid: 11226344
references.citation: Am Nat. 1998 Jul;152(1):24-44
references.pmid: 18811399
references.citation: Proc Natl Acad Sci U S A. 2007 Aug 14;104(33):13219-24
references.pmid: 17686992
references.citation: Science. 1999 Apr 2;284(5411):65-71
references.pmid: 10102822
references.citation: N Engl J Med. 2008 Jul 3;359(1):61-73
references.pmid: 18596274
references.citation: Hum Reprod. 2002 Sep;17(9):2474-9
references.pmid: 12202444
references.citation: Proc R Soc Lond B Biol Sci. 1979 Sep 21;205(1161):531-46
references.pmid: 42059
references.citation: Biol Reprod. 2007 Aug;77(2):247-51
references.pmid: 17429014
references.citation: Proc Biol Sci. 2009 May 22;276(1663):1837-44
references.pmid: 19324786
references.citation: Acta Anat (Basel). 1952;14(1-2):108-23
references.pmid: 14932631
references.citation: Hum Nat. 2002 Jun;13(2):199-238
references.pmid: 26192758
references.citation: J Hum Evol. 2001 May;40(5):437-50
references.pmid: 11322804
references.citation: Science. 2004 Sep 17;305(5691):1736-9
references.pmid: 15375259
references.citation: Am J Phys Anthropol. 1979 Mar;50(3):291-300
references.pmid: 218459
references.citation: Lancet. 1986 May 10;1(8489):1077-81
references.pmid: 2871345
references.citation: Am J Hum Biol. 2003 May-Jun;15(3):380-400
references.pmid: 12704714
references.citation: J Hum Evol. 2002 Dec;43(6):831-72
references.pmid: 12473486
references.citation: Science. 1960 Jul 1;132(3418):14-21
references.pmid: 13835176
references.citation: Am J Epidemiol. 2003 May 15;157(10):923-9
references.pmid: 12746245
references.citation: Am J Hum Biol. 2002 Mar-Apr;14(2):184-205
references.pmid: 11891933
references.citation: J Theor Biol. 2001 Dec 21;213(4):527-45
references.pmid: 11742523
references.citation: Menopause. 2007 Jul-Aug;14(4):668-79
references.pmid: 17549040
references.citation: Am J Hum Biol. 2009 Jan-Feb;21(1):2-15
references.pmid: 18925573
references.citation: J Gerontol A Biol Sci Med Sci. 2007 Jul;62(7):722-7
references.pmid: 17634318
references.citation: Ageing Res Rev. 2006 Feb;5(1):14-32
references.pmid: 16085467
references.citation: J Clin Endocrinol Metab. 1987 Dec;65(6):1231-7
references.pmid: 3119654
references.citation: Am J Hum Biol. 2003 Mar-Apr;15(2):196-208
references.pmid: 12621608
references.citation: Curr Biol. 2007 Dec 18;17(24):2150-6
references.pmid: 18083515
references.citation: Behav Brain Sci. 2005 Oct;28(5):675-91; discussion 691-735
references.pmid: 16262930
references.citation: Am J Primatol. 1995;37(1):25-37
references.pmid: 32005046
references.citation: Hum Reprod. 1996 Jul;11(7):1484-6
references.pmid: 8671489
references.citation: Science. 2002 May 10;296(5570):1029-31
references.pmid: 12004104
references.citation: Proc R Soc Lond B Biol Sci. 1963 Oct 22;158:417-33
references.pmid: 14070052
references.citation: PLoS Genet. 2007 Feb 23;3(2):e24
references.pmid: 17319745
references.citation: Proc Natl Acad Sci U S A. 1991 Feb 15;88(4):1134-7
references.pmid: 1996315
references.citation: Biol Reprod. 2008 Sep;79(3):398-406
references.pmid: 18495681
references.citation: Mol Biol Evol. 2003 Mar;20(3):424-34
references.pmid: 12644563
references.citation: Soc Biol. 2002 Fall-Winter;49(3-4):185-205
references.pmid: 14652917
references.citation: Am J Epidemiol. 2002 Oct 1;156(7):627-33
references.pmid: 12244031
references.citation: Mech Ageing Dev. 2007 May-Jun;128(5-6):355-64
references.pmid: 17459455
references.citation: J Gerontol A Biol Sci Med Sci. 2000 Jun;55(6):B264-73
references.pmid: 10843342
references.citation: J Hum Evol. 1999 May;36(5):461-85
references.pmid: 10222165
references.citation: Endocr Rev. 2009 Aug;30(5):465-93
references.pmid: 19589949
references.citation: Q Rev Biol. 2004 Mar;79(1):3-50
references.pmid: 15101252
references.citation: Demography. 1979 Aug;16(3):439-54
references.pmid: 510638
references.citation: J Gerontol A Biol Sci Med Sci. 2009 Jul;64(7):740-4
references.pmid: 19414513
references.citation: J Am Geriatr Soc. 2005 Dec;53(12):2184-9
references.pmid: 16398907
references.citation: Anim Behav. 2000 Apr;59(4):895-896
references.pmid: 10792945
references.citation: Oxf Rev Reprod Biol. 1989;11:61-109
references.pmid: 2697833
references.citation: Proc Natl Acad Sci U S A. 2010 Jan 26;107 Suppl 1:1718-24
references.pmid: 19966301
references.citation: Evolution. 2007 Jun;61(6):1265-76
references.pmid: 17542838
references.citation: Hum Biol. 1959 Feb;31(1):54-73
references.pmid: 13640652
references.citation: Biol Reprod. 2008 Sep;79(3):407-12
references.pmid: 18495682
references.citation: Am J Phys Anthropol. 2000 Nov;113(3):435-50
references.pmid: 11042542
references.citation: Maturitas. 1981 Aug;3(2):157-66
references.pmid: 6793812
references.citation: Proc Natl Acad Sci U S A. 2008 May 20;105(20):7124-8
references.pmid: 18458325
references.citation: Am J Primatol. 2008 Aug;70(8):766-77
references.pmid: 18506732
references.citation: Ann N Y Acad Sci. 2010 Aug;1204:43-53
references.pmid: 20738274
references.citation: J Hum Evol. 2007 Apr;52(4):443-54
references.pmid: 17289113
references.citation: Proc Natl Acad Sci U S A. 1998 Feb 3;95(3):1336-9
references.pmid: 9448332