Grandmother hypothesis
The grandmother hypothesis is a theory to explain why menopause, rare in mammal species, arose in human evolution and how a long post-fertile period (up to one third of a female's life-span)[1] could confer an evolutionary advantage.
Menopause should, by Darwinian Theory, reduce fecundity, making menopause intriguing from an evolutionary perspective, as natural selection will operate vigorously to delay the expression of harmful alleles.[2] It is, therefore, advocated that there is a genetic ‘intention’ for postmenopausal females.
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[edit] Background
In female placentals, the number of ovarian oocytes is fixed during embryonic development; possibly as an adaptation to reduce the accumulation of mutations.[3] At birth there are, typically, one million ova; when menopause begins, only 400 eggs would have actually matured.[4] The intriguing question is why somatic cells decline at a slower rate and why humans invest more in somatic longevity than other primates.[5]
More important than the question of why longevity has been extended, however, is why selection has not adjusted female life-history to match; as it appears to have done with males. The most frequently cited adaptive causes for the menopause are variations on the ‘mother’, or ‘grandmother’ hypothesis. These theories advocate that the high costs attributed to female reproduction could prevail over the benefits of continuous propagation. It is true that with advancing age and decreasing fertility, there is also a corresponding increase of miscarriages and birth defects, such as Down’s syndrome.[6] Age is less significant in the increased foetal abnormalities than is the number of the ova left in the ovarian follicular reserves.[7]
A possible explanation is the rate of oocyte depletion. With ova numbers fixed, it is logical to think extension of fertility would require increasing the size of the ovaries, which may be a limiting factor. However, the rate of atresia accelerates before age 40; if it did not, oocyte depletion would not occur until age 70.[8] Regardless, limited oocyte production and accelerated follicular atresia only explains why fertility ends in females and not why they survive for many years in a post-reproductive state.
See also non-adaptive hypotheses for menopause
[edit] The grandmother effect
G.C. Williams was the first to posit[9] that menopause might be an adaptation. Williams suggested that at some point during evolution, it became advantageous for females to stop "dividing [their] declining faculties between the care of extant offspring and the production of new ones" (p. 408). Since a female's dependent offspring would die as soon as she did, he argued, older mothers should stop producing new babies and focus on the offspring they already had. In so doing, they would avoid the risk of dying during childbirth and thereby eliminate a potential threat to the continued survival of current offspring.
In addition, postmenopausal women can contribute knowledge and skills to other group members to enhance group fitness.[10] If the other group member receiving investment were kin, then this would increase the fitness of a post-menopausal woman.[2][11]
This kin selection emerged with climate-driven changes, around 1.8 –1.7 million years ago, in female foraging and food sharing practices. These adjustments increased juvenile dependency, forcing mothers to opt for a low-ranked, common food source (tubers) that required adult skill to harvest and process. Such demands constrained female birth intervals and consequently their fertility; thus providing an opportunity for selection to favour the grandmother hypothesis.[12]
[edit] Evidence for
If the grandmother effect were true, post-menopausal women should work harder than their still-fertile counter-parts and share those proceeds with their kin. Studies of Hadza women have provided such evidence,[13] but they have not shown that this behaviour actually increased the grandmother’s fitness by producing more grandchildren. Furthermore, the role of Hadza men, who contribute 96% of the mean daily intake of protein, was ignored.[14] Other studies also demonstrated reservations about behavioural similarities between the Hadza and our ancestors.[15]
Quantitative models, however, showed increased fitness, but not significantly.[16] One of the reasons given for low benefits is the inflexibility of the age at which menopause occurs. It could be that fertility behaviour is adjusted depending on long-term expected fitness.[17] Such choice, however, is not available to an already infertile female – she cannot ‘choose’ to redirect investment.
[edit] The grandmother effect and longevity
It is said that the grandmother hypothesis “is a central determinant of our longevity”;[18] yet available data suggests that the reverse may be so – our longevity is a central determinant of the grandmother hypothesis.[17] Further analysis of the historical data found that the length of a female’s post-reproductive lifespan was reflected in the reproductive success of her offspring and the survival of her grandchildren.[19]
[edit] Maternal vs paternal grandmothers
Similar studies found comparative affects but only in the maternal grandmother – paternal grandmothers had a detrimental effect on infant mortality,[20][21] as well as differing assistance strategies for maternal and paternal grandmothers. Maternal grandmothers concentrate on offspring survival, whereas paternal grandmothers increase birth rates.[22] These finding are actually consistent with the grandmother hypothesis because of paternity uncertainty. Equally, a grandmother could be both a maternal and paternal grandmother and thus in division of resources, a daughter’s offspring should be favoured.
[edit] Evidence against
Such historical studies are, however, unable to quantify grandmotherly assistance; they are merely correlations between infant mortality and the existence of a grandparent. Those studies that have calculated assistance to both offspring and grandchildren have not found appreciable affects to warrant termination of fertility as early as 50.[16]
Another problem concerning the grandmother hypothesis is that it requires a history of female philopatry and yet present day evidence shows that the majority of hunter-gatherer societies are patriarchal.[23] Others dispute the hypothesis, arguing that the grandmother herself will use up resources that could be used for new young.
In addition, all variations on the mother, or grandmother effect, fail to explain longevity with continued spermatogenesis in males. It also fails to explain the detrimental effects of losing ovarian follicular activity, such as osteoporosis, osteoarthritis, Alzheimer’s disease and coronary artery disease.[24]
Alternatively, the debilitating symptoms that usually accompany menopause in Western cultures could be seen as a natural cull of non-reproductive members of a species. Hot flashes, loss of short term memory, decreased ability to concentrate and difficulty in the learning of new tasks would, in the wild, leave the sufferer at greater risk from predators and topographical dangers such as falls from a height. This natural cull would leave more food, usually in relatively short supply, for the reproductive members of a species whose youth could mean that they are less experienced at finding it. However, cross-cultural studies of menopause have found that menopausal symptoms are quite variable among different populations, and that some populations of females do not recognize, and may not even experience, these "symptoms".[25] This high level of variability in menopausal symptoms across populations brings into question the plausibility of menopause as a sort of "culling agent" to eliminate non-reproductive females from competition with younger, fertile members of the species.
[edit] Conclusion
The grandmother effect needs to establish that longevity, menopause expression, and redirected investment commonly occurred in our ancestral past. Evidence from contemporary hunter-gather societies and historical data produce mixed results. Many theoretical assumptions, such as differing strategies for maternal and parental grandparents, have been shown. The fundamental flaw, however, is the exclusion of male assistance and benefits from continued fertility.[26]
[edit] References
- ^ Harman, S.M.; Talbert, G.B. (1985). "Reproductive aging". In C. E. Finch and L. Hayflick. Handbook of the Biology of Aging. Van Nostrand Reinhold. pp. 457–510.
- ^ a b Dawkins, Richard (1976). The selfish gene. Oxford: Oxford University Press. ISBN 1840462388.
- ^ Ellison, P.T. (2001). On Fertile Ground: Ecology, Evolution and Human Reproduction. Harvard University Press.
- ^ Pizzorno, L.; Pizzorno Jr., J.E and Murray, M. (2002). Menopause. Natural Medicine Instructions for Patients. Elsevier Science Ltd. pp. 209–214.
- ^ Hawkes, K. (2003). "Grandmothers and the evolution of human longevity". American Journal of Human Biology 15 (3): 380–400. doi:10.1002/ajhb.10156. PMID 12704714.
- ^ Finch, C.E. (1990). Longevity senescence and the genome. London: University of Chicago Press. ISBN 0226248887.
- ^ Brook, J.D.; Gosden, R.G. and Chandley, A.C. (1984). "Maternal Aging and Aneuploid Embryos: Evidence from the Mouse that Biological and not Chronological Age is the Important Influence". Human Genetics 66 (1): 41–45. doi:10.1007/BF00275184. PMID 6538182.
- ^ Gosden, R.G.; Faddy, M.J. (1998). "Ovarian aging, follicular depletion and steroidogenesis". Experimental Gerontology 29 (3–4): 265–274. doi:10.1016/0531-5565(94)90006-X. PMID 7925747.
- ^ Williams, GC (1957). "Pleiotropy, natural selection, and the evolution of senescence". Evolution 11 (4): 398–411. doi:10.2307/2406060. JSTOR 2406060.
- ^ Campbell, B. (1971). Human Evolution: An Introduction to Man’s Adaptations. Aldine. ISBN 0202020134.
- ^ Alexander, R.D. (1974). "The Evolution of Social Behaviour". Annual Review of Ecology and Systematics 5: 325–83. doi:10.1146/annurev.es.05.110174.001545.
- ^ O’Connell, J.F.; Hawkes, K., Blurton Jones, N.G. (1999). "Grandmothering and the evolution of Homo erectus". Journal of Human Evolution 36 (5): 461–485. doi:10.1006/jhev.1998.0285. PMID 10222165.
- ^ Hawkes, K.; O’Connell, J.F. and Blurton Jones, N.G. (1997). "Hazda Women's Time Allocation, Offspring Provisioning, and the Evolution of Long Postmenopausal Life Spans". Current Anthropology 38 (4): 551–577. doi:10.1086/204646.
- ^ Gurven, M.; Hill, K. (1997). "Comments on 'Hazda Women's Time Allocation, Offspring Provisioning, and the Evolution of Long Postmenopausal Life Spans'". Current Anthropology 38 (4): 566–567.
- ^ Gibbons, A. (1997). "Why Life After Menopause?". Science 276 (5312): 535. doi:10.1126/science.276.5312.535b.
- ^ a b Hill, K.; Hurtago, A.M. (1996). Ache Life History: The Ecology and Demography of a Foraging People. New York: Hawthorne. ISBN 0202020371.
- ^ a b Winterhalder, B.; Smith, E.A. (2000). "Analysing Adaptive Strategies: Human Behavioural Ecology at 25". Evolutionary Anthropology 9 (2): 51–72. doi:10.1002/(SICI)1520-6505(2000)9:2<51::AID-EVAN1>3.0.CO;2-7.
- ^ Hawkes,, K. (2004). "Human longevity: The grandmother effect". Nature 428 (6979): 128–129. doi:10.1038/428128a. PMID 15014476.
- ^ Lahdenperä, M.; Lummaa, V., Helle, S., Tremblay, M. & Russell, A. F. (2004). "Fitness benefits of prolonged post-reproductive lifespan in women". Nature 428 (6979): 178–181. doi:10.1038/nature02367. PMID 15014499.
- ^ Voland, E.; Beise, J. (2002). "Opposite Effects of Maternal and Paternal Grandmothers on Infant Survival in Historical Krummörn". Mpidr Wp 2001-026.
- ^ Jamison, C.S.; Cornell L.L., Jamison P.L. & Nakazato H. (2002). "Are all grandmothers equal? A review and a preliminary test of the grandmother hypothesis in Tokugawa Japan". American Journal of Physical Anthropology 119 (1): 67–76. doi:10.1002/ajpa.10070. PMID 12209574.
- ^ Mace, R.; Sear, R. (2004). "Are Humans Communal Breeders?". In Voland, E., Chasiotis, A. and Schiefenhoevel, W.. Grandmotherhood – the Evolutionary Significance of the Second Half of Female Life. Rutgers University Press.
- ^ Peccei, J. S. (2001). "A critique of the grandmother hypotheses: Old and new". American Journal of Human Biology 13 (4): 434–452. doi:10.1002/ajhb.1076. PMID 11400215.
- ^ Massart, F.; Reginster, J.Y. and Brandi, M.L. (2001). "Genetics of Menopause-Associatred Diseases". Maturitas 40 (2): 103–116. doi:10.1016/S0378-5122(01)00283-3. PMID 11716989.
- ^ Melby, Melissa K. (2005). "Vasomotor symptom prevalence and language of menopause in Japan". Menopause 12 (3): 250–257. doi:10.1097/01.GME.0000146108.27840.D9?. PMID 15879913.
- ^ Hudson, M. (2005). Why do females (especially human females) often survive beyond an age at which they can reproduce?. Brighton: Sussex University Press.
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