Grandmother hypothesis

The grandmother hypothesis is a hypothesis to explain the existence of menopause in human life history by identifying the adaptive value of extended kin networking. It builds on the previously postulated " mother hypothesis" which states that as mothers age, the costs of reproducing become greater, and energy devoted to those activities would be better spent helping her offspring in their reproductive efforts. It suggests that by redirecting their energy onto those of their offspring, grandmothers can better ensure the survival of their genes through younger generations. By providing sustenance and support to their kin, grandmothers not only ensure that their genetic interests are met, but they also enhance their social networks which could translate into better immediate resource acquisition. This effect could extend past kin into larger community networks and benefit wider group fitness.

Background

One explanation to this was presented by G.C. Williams who was the first to posit that menopause might be an adaptation. Williams suggested that at some point it became more advantageous for women to redirect reproductive efforts into increased support of existing offspring. Since a female's dependent offspring would die as soon as she did, he argued, older mothers should stop producing new offspring and focus on those existing. In so doing, they would avoid the age-related risks associated with reproduction and thereby eliminate a potential threat to the continued survival of current offspring. The evolutionary reasoning behind this is driven by related theories.

Kin selection

Kin selection provides the framework for an adaptive strategy by which altruistic behavior is bestowed on closely related individuals because easily identifiable markers exist to indicate them as likely to reciprocate. Kin selection is implicit in theories regarding the successful propagation of genetic material through reproduction, as helping an individual more likely to share one's genetic material would better ensure the survival of at least a portion of it. Hamilton's rule suggests that individuals preferentially help those more related to them when costs to themselves are minimal. This is modeled mathematically as $rb > c$. Grandmothers would, therefore, be expected to forgo their own reproduction once the benefits of helping those individuals (''b'') multiplied by the relatedness to that individual (''r'') outweighed the costs of the grandmother not reproducing (''c'').

Evidence of kin selection emerged as correlated 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 IBIs thus providing an opportunity for selection to favor the grandmother hypothesis.

Parental investment

Parental investment, originally put forth by Robert Trivers, is defined as any benefit a parent confers on an offspring at a cost to its ability to invest elsewhere. This theory serves to explain the dynamic sex difference in investment toward offspring observed in most species. It is evident first in gamete size, as eggs are larger and far more energetically expensive than sperm. Females are also much more sure of their genetic relationship with their offspring, as birth serves as a very reliable marker of relatedness. This paternity uncertainty that males experience makes them less likely than females to invest, since it would be costly for males to provide sustenance to another male's offspring. This translates into the grandparental generation, as grandmothers should be much more likely than grandfathers to invest in the offspring of their children, and more so in the offspring of their daughters than sons.

The grandmother effect

Evolutionary theory dictates that all organisms invest heavily in reproduction in order to replicate their genes. According to parental investment, human females will invest heavily in their young because the number of mating opportunities available to them and how many offspring they are able to produce in a given amount of time is fixed by the biology of their sex. This inter birth interval (IBI) is a limiting factor in how many children a woman can have because of the extended developmental period that human children experience. Extended childhood, like the extended post-reproductive lifespan for females, is unique to humans. Because of this correlation, human grandmothers are well-poised to provide supplemental parental care to their offspring's children. Since their grandchildren still carry a portion of their genes, it is still in the grandmother’s genetic interest to ensure those children survive to reproduction.

Reproductive senescence

The mismatch between the rates of degradation of somatic cells versus gametes in human females provides an unsolved paradox. Somatic cells decline more slowly, and humans invest more in somatic longevity relative to other species. Since natural selection has a much stronger influence on younger generations, deleterious mutations