The origin of menopause has puzzled evolutionary biologists for the last half-century. Three new studies attempt illumination. The real question, though, is probably not: Why menopause? Rather, it is: Why do women long outlive their fertility?



Human ovaries tend to shut down by age 50 or even younger, yet women commonly live on healthily for decades. This flies in the face of evolutionary theory that losing fertility should be the end of the line, because once breeding stops, evolution can no longer select for genes that promote survival.



The most popular explanation, the "grandmother hypothesis," argues that a generous post-reproductive life span makes sense if a grandmother improves the survival and reproduction of her grandchildren, thus ensuring continuation of her own genes—including genes that contribute to longevity. But skeptics say the math is askew. From an evolutionary perspective, it is hardly ever better for a woman to give up a chance to bear additional children of her own, and so pass on half her genes, for the sake of improving the survival of her grandchildren, who carry only a quarter of her genes.



"The problem is that these grandmother benefits aren't big enough to ever favor stopping breeding between the ages of 40 and 50," says Michael Cant, an evolutionary biologist at the University of Exeter in England and co-author of a new study on the genesis of menopause published this week in Proceedings of the National Academy of Sciences USA. "When you look at data from hunter-gatherers and other natural fertility populations, the sums just don't add up." Grandmothers do benefit their descendants, he says, but the genetic payoff is small compared with those of producing another child.



Cant and co-author Rufus Johnstone, an evolutionary biologist at the University of Cambridge in England, used game theory to argue that menopause is early cessation of reproduction that originated through reproductive conflict between generations. In most cooperatively breeding species, reproduction is suppressed in younger females, who act as helpers to older reproducing females. By contrast, they say, younger women in human social groups win the reproductive sweepstakes, because the older ones stop having babies.



"We showed that, compared to other primates that exhibit a post-reproductive life span, humans really stand out, because there is absolutely no overlap in reproduction between generations," Cant says. "Women stop breeding on average when the next generation starts to breed."



This makes evolutionary sense, Cant and Johnstone say, because, contrary to most mammals, young women tend to move to their mates' communities, where they become immigrants whose only genetic kin are their own children. There is no genetic profit in helping their mothers-in-law bear more children, because they will not share any genes with those children. But an older woman who helps her son's wife reproduce will benefit by bequeathing 25 percent of her genes to her grandchildren.



"We show that the mother-in-law's best strategy is to stop breeding, avoid competition, and allow the daughter-in-law to breed and help her," Cant says. "It's the first time anyone has taken the idea that humans evolved with this sex bias in dispersal and looked at the implications for how these conflicts will be resolved within the family."



The mother of the grandmother hypothesis, anthropologist Kirsten Hawkes of the University of Utah, says Cant and Johnstone are right to focus on intergenerational conflict. Elephants have babies in their 60s, and some whales give birth in their 80s. "It's clearly something selection can adjust," she says. "So explaining why it hasn't in us has to be part of the story." But she disputes their claim that female-bias dispersal is, in fact, the universal human/ape residence pattern, pointing out that half of the young female chimps at anthropologist Jane Goodall's Gombe Stream Research Center remain with their mothers, and that recent studies show that hunter-gatherers often live with the wife's family as well.



Another explanation for menopause is the "mother hypothesis," which holds that it occurs because older mothers might profit more, genetically speaking, by investing resources in their existing children than in giving birth to new ones. Researchers at the Max Planck Institute for Demographic Research in Rostock, Germany, make the case for this in the American Journal of Physical Anthropology (AJPA), concluding that menopause is advantageous when a woman has aged enough to face an increased risk of stillbirth, birth defects and her own death in childbirth.



Researchers of a different AJPA study, based on 400 years' worth of data on births in Costa Rica, believe that postmenopausal longevity is associated with an increased number of children but a decreased number of grandchildren—a finding that supports mothers over grandmothers.



"We're not saying grandmothers do not provide benefits in some societies," says study co-author Lorena Madrigal, an anthropologist at the University of South Florida in Tampa. But, "we should not assume that one pattern fits all."



Data on great ape fertility is spotty, but what there is shows that our closest relatives—chimps, bonobos, gorillas, even orangutans—stop having babies about the same age that we do: the late 30s. The difference is, they generally die a short time later. "The thing that makes us different from apes is not the age of fertility decline, it's the lack of aging in other systems," says Hawkes. "I have been saying this for a long time and I don't think it's what anybody is hearing. Probably what a lot of people are prepared to listen to is the way Cant and Johnstone have framed this, that the real question is: 'Why do we stop [reproducing] so early?' I think the bottom line is that, compared with our closest living relatives, we don't."