People often ask me whether natural selection continues to operate on modern humans in industrialised societies, even though technology has liberated so many from hunger and early death. My answer is always an unambiguous “Yes!”.
A recently published paper illustrates a dramatic episode of selection that happened in China a mere 50 years ago, the effects of which continue to reverberate through Chinese society. It’s an example that further illustrates how selection in the sex ratio is always happening, even in the most privileged modern societies.
A dramatic graph (see below) from the paper caught my attention. It shows the sex ratio of babies to women in China between 1938 and 1982. The data came from a massive retrospective survey of the fertility of 300,000 women.

What staggered me, as it did Shige Song who wrote the paper in which the graph appeared, was the massive drop in the proportion of male births. Something happened in China in the early 1960s that massively changed the sex ratio.
That “something” was a famine – probably history’s largest. Between the end of 1958 and the start of 1962, the ill-conceived economic initiatives of the Great Leap Forward led to a China-wide famine that killed 20-30 million people.
The famine also caused a precipitous drop in the number of births (see figure below), as women of child-bearing age starved and were unable to conceive or carry their foetuses to term.

These graphs imply a human tragedy so immense it defies my ability to describe. But they also illustrate an important aspect of evolutionary biology that remains relevant even in the most modernised societies.
Sex ratio and selection
The ratio of males to females in a population, including the human population, is always under selection. That’s because if one sex becomes rare, that sex experiences less competition come mating time. So parents who can bias conception – or even the care they invest in their young – toward that rarer sex will, in time, have more grandchildren.
In most species, this effect of competition keeps the sex ratio close to even. But that doesn’t mean every family should benefit equally via sons and daughters.
For one thing, in most mammals it takes more effort to raise a successful son who will go on to hold a territory and be a successful breeder. But raising such a successful son is like winning the evolutionary lottery.
That’s why a red deer doe in good condition is more likely to give birth to a son – who has a good chance of growing big and winning dominion over a harem – than a daughter. Whereas mums-to-be who are in good enough condition to carry and care for a fawn, but not in the peak condition likely to yield a future king of the forest, tend to have daughters.
This is the Trivers-Willard effect, one of the most original and powerful ideas to emerge from modern evolutionary biology. It has been confirmed in hundreds of studies from wasps (where more care makes better females) to horses.
In mammals, it seems conditions in the womb might affect the survival of male or female embryos. Certainly when in-vitro fertilised cattle embryos are reared in a glucose-rich medium, only males survive, but female embryos thrive on less rich media.
Trivers and Willard actually came up with their theory by thinking about humans. Trivers mentioned to a class he was teaching that girls often marry upward into families of higher status and greater wealth. When this happens, there are too many girls competing for the wealthiest boys. And too many poor boys competing for the few poor girls who haven’t already married up.
Willard – a student in the class – suggested wealthy families might benefit from biasing conception or investment toward sons, and that poorer families should do the opposite. Crucially, the effects of too many boys born into wealthy families and excess girls in poorer families would balance out, leaving the overall sex ratio approximately equal.
Evidence suggests this is exactly what happens. Some 60% of the children born to billionaire families are sons. If that ratio was repeated Australia-wide, men and boys would outnumber women and girls by about four million.
Junior wives in polygynous Rwandan marriages suffer low status and often agonising poverty, and they have overwhelmingly more daughters than higher-status first wives or the wives of monogamously married men.
Trivers-Willard effects manifest after children are born too. In the pre-industrial German parish of Leezen, sons in wealthy land-owning families were far more likely than their sisters to survive to their first birthday, whereas the opposite was true in poorer families. That’s probably due to subtle, even sub-conscious, differences in how parents fed and cared for their sons and daughters.
The Great Leap Forward
If the mechanisms underlying Trivers-Willard effects are aligned – as we suspect – with nutritional conditions, then we might expect dramatic sex ratio fluctuations when a whole population experiences extreme food shortage.
Yet studies of two of the 20th century’s most dramatic famines – the Dutch Hunger Winter of 1944-5 and the 1942 Leningrad Siege – showed equivocal associations between famine and sex ratios. Crucially, these famines lasted seven and six months respectively – enough to kill vast numbers of people but perhaps not long enough to bias the birth of an entire cohort.
It’s also possible that the least-starved mothers, deprived as they were, produced sons because they were in relatively better condition than the other mothers.
Trivers-Willard effects often seem to be relative. Only the very wealthiest and highest status families experience the strong bias toward having more sons, irrespective of their absolute level of wealth. That would explain how billionaires could have dramatically more sons than mere multi-millionaires. And that might explain why the sex ratio doesn’t bounce around too dramatically from year to year, responding to lean and fat years.
But Song’s paper, published last week, shows that a long-lasting, severe famine can change the sex ratio at birth. Both birth rate and sex ratio headed south about a year into the famine and remained there until about two years after the famine ended.
It seems that for the Trivers-Willard effect to bias the sex ratio of an entire cohort, the famine must last for some time before the onset of pregnancy.
And presumably the drop in birth rate comes from male embryos and foetuses faring worse than females during famine, either failing to implant, or miscarrying at higher rates.
China’s sex ratios are notoriously male-biased and becoming more so, but the worst famine in history created one tiny cohort in which almost as many girls were born as boys.
The graphs above are republished courtesy of the Royal Society London and originally appeared in Does famine influence sex ratio at birth? Evidence from the 1959-1961 Great Leap Forward Famine in China, by Shige Song, published online before print March 28, 2012, doi: 10.1098/rspb.2012.0320.
Gideon Polya
Sessional Lecturer in Biochemistry for Agricultural Science at La Trobe University
Excellent article. This finding of a slightly but significantly increased male to female sex ratio at birth with increased prosperity and satiety is a useful bias in relation to the current urgent linked problems of tackling human overpopulation and the worsening climate crisis (see "2011 Climate Change Course": https://sites.google.com/site/300orgsite/2011-climate-change-course ). Thus only females can have children and from a simple animal husbandry perspective - and setting aside culture and gender…
Read moreDale Bloom
Analyst
I would think males are very important. They are the ones who mostly build the dams and develop the crops to avoid starvation.
Without males, famines would likely increase in frequency and duration.
Mal O'Keeffe
Agricultural Scientist
In China, women constitute about 70 percent of the agricultural labour force and perform more than 70 percent of farm labour. The general pattern is - the poorer the area, the higher women's contribution.
The world over, 70% of the agricultural workers, 80% of food producers, and 10% of those who process basic foodstuffs are women and they also undertake 60 to 90% of the rural marketing; thus making up more than two-third of the workforce in agricultural production. Despite the fact that women produce much of the food in the developing world, they also remain more malnourished than most men.
I could go on with more information on the vital role that women play in feeding the world but I know from previous comments that you do not argue in good faith, or with any discernible knowledge but only with the sole purpose to repetitively express your contempt for women.
Dale Bloom
Analyst
Mal,
I have not expressed contempt for women. I think you've made that up.
John Holmes
Agronomist - semi retired consultant
Some FAO data years old showed that it is the women who tend to do most of the hand weeding of crops where mechanical or herbicidal weed control is not used. The studies looked also at the energy requirements as well. Hand weeding does use not insignificant amounts of energy. Sure it is not fueled directly by fossil fuel.
For those who wish to have an "organic" existence, there can be major limitations in production as there can be serious limitations in work force availability to keep the…
Read moreLisa Milne
Education at Southern Cross University
Hi Mal,
Use the report abuse button - much more effective (as you know, but it is tempting I know) than engaging with Dale the troll.
Cheers
Gordon Young
Environmental Consultant
Fascinating phenomena, but I'm confused as to how the Trivers-Willard effect functions in higher value societies.
I can accept that within poorer communities nutrition and selective favourism/neglect of children may lead to richer families having more sons, and poorer families more daughters, due to some sort of biological programming. It also makes a lot of sense with animals (and interesting that bees reverse the gender bias seen in mammals!).
But this doesn't hold when comparing billionaire…
Read moreRob Brooks
Rob Brooks is a Friend of The Conversation.
Professor of Evolutionary Ecology; Director, Evolution & Ecology Research Centre at University of New South Wales
I'm intrigued, too, about the effect of relative wealth on sex ratios. Your comment made me realise that nobody has looked at sex ratios in 'mere' multi-millionaires and that I'm giving the wrong impression that they aren't male biased where, really, no good evidence exists. What is robust is that billionaires have more sons AND that they gain more grandchildren per son than they do per daughter. But being on Forbes' billionaires list was really just a convenient way to circumscribe the sample…
Read moreDale Bloom
Analyst
Rob Brooks
The only usefulness in any of this might be regards IVF, and IVF should be banned for a number of reasons.
If a man is selected to be the father of a child by selecting him from a catalogue that contains his picture and a small bio, then I would think IVF must be one of the most degenerate technologies.
But you overlooked that part.
Gordon Young
Environmental Consultant
Thank you for the explanation Rob, that was very helpful. I think I have a better understanding of the effect and its application in developed populations.
I look forward to hearing more as the research and our understanding develops!
Yoron Hamber
Thinking
Nah, you're drawing too big a conclusion from too little statistics there me thinks :) The difference in living between being a mere millionaire and a billionaire I doubt to differ that much. They should both have the means to train and eat correctly. so that effect needs a lot more proofs than this. And you can also use the statistics from Countries as Sweden and compare it to what we eat to check your idea. We are big on statistics here in the Nordic countries, been doing it for quite some time. It can be a very powerful tool for finding trends and connections..
But your conclusions on the famines effects are truly fascinating and something I've never seen before, and, there you also have good statistics as it seems to me? So yes, maybe there is just such a phenomena coupled to famines.
Sean Lamb
Science Denier
Bit of mixed bag this article. The change in sex ratio in the Chinese famine is interesting, but the effect is relatively minor (exaggerated by the choice of scale on the y axis) - .522 to .510. Since we know that male infant mortality is significantly higher than female, is it so surprising that in a period of extreme strain this would extend backwards into the gestation period? And that is assuming perfect data collection and that rural areas didn't tend to not record births where the infant…
Read moreRob Brooks
Rob Brooks is a Friend of The Conversation.
Professor of Evolutionary Ecology; Director, Evolution & Ecology Research Centre at University of New South Wales
Rwandan wives and Forbes billionaires are what authors call "examples". A retrospective sample of 300,000 Chinese women is hardly a trivial piece of empirical evidence.
While Trivers-Willard effects don't always seem to occur, and they aren't consistent there are literally thousands of tests, of which hundreds have been performed using big samples in humans. Here's one - the first one that fell to hand when I searched - http://rspb.royalsocietypublishing.org/content/274/1624/2491.short
These studies are about small deviations in sex ratio - deviations that are only detectible in huge samples. There are indeed sex biases in Australian Statistical Local Areas, but those are confounded by the fact that people move to neighbourhoods where the best schools are - so families with only boys might be found in neighbourhoods where the best boys' schools are and vice versa for families with only girls.
Dale Bloom
Analyst
I would agree,
The figures for how many people died range from 20 to 30 million, and if it is not accurately known how many people died during this time, it is difficult to accept that accurate figures were kept of how many babies were born.
When it comes to issues of gender, very little from university academics can be believed.
Sean Lamb
Science Denier
Well then use census data or occupation - eg doctors versus hospital cleaners. Forbes billionaires are hardly a large or typical sample. If by Trivers-Willard you mean simply that male infant mortality is traditionally higher and higher income, higher status families have lower infant mortality, then you subvert the purported evolutionary benefit as the driver of it.
If you are looking for evolutionary explanation you need to
Read morea) show that there exists or could exist a molecular/genetic mechanisms…
Rob Brooks
Rob Brooks is a Friend of The Conversation.
Professor of Evolutionary Ecology; Director, Evolution & Ecology Research Centre at University of New South Wales
Research into mechanisms is not an alternative to evolutionary/adaptive explanation. The two types of explanation are complementary aspects of science. A good theoretic prediction that is upheld without a confirmed mechanism is still good science.
My report is on a paper that was peer-reviewed and published in the worlds oldest and one of its most prestigious journals. From my reading of the paper I find the story convincing and I hope you read it and decide if the evidence holds up rather than…
Read moreSean Lamb
Science Denier
Oh dear, where to start.
First passing over the argument from authority "peer reviewed journal", peer reviewed journals are filled with work that can't be reproduced and/or are completely contradicted a few years later. In point of fact pubmed only turns up 44 papers on Trivers-Willard, suggesting it is not a particularly popular concept either with demographers or evolutionists (and of that 44 by no means all support the model - lactating dairy cows being a notable holdout in this prestigious…
Read moreRob Brooks
Rob Brooks is a Friend of The Conversation.
Professor of Evolutionary Ecology; Director, Evolution & Ecology Research Centre at University of New South Wales
Sean,
I'm always happy to discuss ideas with anyone who seems willing to have an open discussion - but not with folks (and there are plenty) who merely wish to provoke or to blather about their half-baked conspiracy theories. My call for you to scrutnise the original paper published in a reputable journal if you find this topic so interesting wasn't an "argument from authority" but an appeal to take the paper on it merits which, in my opinion, it deserves.
Google Scholar (the only tool available…
Read moreDale Bloom
Analyst
So you are saying boys take more resources to raise than girls. I wonder if anyone has ever asked human parents about that, or just made up stories.
“That’s why a red deer doe in good condition is more likely to give birth to a son – who has a good chance of growing big and winning dominion over a harem – than a daughter”
Comparing humans to other species I would regard as very subjective and suspect science. I have noticed that when humans are compared to other species, the other species is always subjectively selected.
Why not compared humans to mutton birds, that annually fly half-way around the world to nest and raise young. Or compare humans to sea turtles, that don’t start to lay eggs until they are 40-50 years old. Or compare humans to praying mantis, that can eat their young.
Sean Lamb
Science Denier
1. You can always access Pubmed so long as you have an internet connection, so it disingenuous to pretend otherwise. I accept that T-W is more widespread that the Pubmed search (which I presume you checked!) suggests. Google Scholar results will be inflated by the fact it checks not just abstracts (like Pubmed) but the entire paper including references. Also older abstracts don't get searched by Pubmed.
2. I am not sure what you mean by conspiracy theories. I worked for a period in the lab…
Read moreSean Lamb
Science Denier
For completeness I will add the rest of the LE Threlkeld data (found on page 283). Data collection ranged from Bateman's Bay to Penrith, to Berrima, to Port Sevens (Threlkeld was himself based at Lake Macquarie)
Sir I herewith enclose the returns for the Aborigines consisting of 102 the number transmitted to me with thanks for the loan of them; but regret that their imperfect state in each year renders it impossible to use them as a criterion to judge of the increase or decrease of the Aborigines, many places not being returned. I have forwarded with them a copy shewing the comparitive [sic] number of the sexes, which is all the available service the returns could afford.
[snip detailed breakdown by district and station]
1838 Men 754, Women 448, Boys 205, Girls 153
1839 Men 737, Women 392, Boys 175, Girls 136
1840 Men 341, Women 208, Boys 130, Girls 100
John Holmes
Agronomist - semi retired consultant
From my limited reading re 'first contact' of two very different civilizations, is/was there not a tendency for women and children to 'go bush' when strange (male & armed?) outsiders appear? If so reported numbers could be quite distorted. A similar more recent situation would possible be the N.G. highlands. I guess the reports/stories etc from anthropologists/ plantation owners/ miners / missionaries / government patrol officers could assist.
Sean Lamb
Science Denier
Those are fair points, another confounding factor would be convicts/settlers/sealers taking sexual partners.
However the gender ratios seem to be found almost as pronounced among children also (however Threlkeld defined child). Moreover Threlkeld seemed to think the gender ratio was real, as he made a point of calculating it and remarking upon it, and he is the man on the spot. At this point, 1830-40, we are beyond first contact and Threlkeld was making every effort to contact the indigenous, learn their language and translate the gospel.
Sean Lamb
Science Denier
Finally on the subject on sex ratios amongst Australian Aborigines, I looked up the classic text Kamilaroi and Kurnai by Fison and Howitt.
Somewhat contrary to my recollection, Fison and Howitt ("periphery anthropologists") were both sceptical of female infanticide but said it was the belief of the then influential "centre athropologists" (ie the UK) John Lubbock and John McLennan. On page 171 to 176 they add some data they got from correspondents based in Fiji.
Navitilevu Hill Tribes (little missionary influence) 1390 Male children to 1144 Female children
Lau Hill tribe (missionary influence) 130 male children to every 100 female children
Wainimala tribes under Wesleyan mission influence
133.66 male children to every 100 female children
Rob Brooks
Rob Brooks is a Friend of The Conversation.
Professor of Evolutionary Ecology; Director, Evolution & Ecology Research Centre at University of New South Wales
Thanks for those toxoplasma references. I'm very pleased to have them and the effects are really quite dramatic. Did you the interesting profile on Flegr's work in The Atlantic a few weeks ago?
I haven't made it back to UNSW yet but will dig out the old references you mention. I shouldn't rule out infanticide purely because it sounds implausible. Given the widespread effects of infanticide in northern India and China (some of which seem to be difficult to comprehend without recourse to Trivers-Willard and other adaptive sex ratio theories) it seems that it's quite a plausible, if offensive, mechanism.
Daniel Barbezat
Professor of economics, Amherst Collegaze
In an extreme famine, overall infant mortality rates rise and overall fertility falls. If some of the sex ratio difference is traditionally due to the infanticide of girls, then maybe fewer female newborns were selectively removed, as having a child at all seemed more difficult. This is a grim conjecture but one that should be addressed.
Dale Bloom
Analyst
Daniel,
I really don’t know how the author missed this, but boy babies are more likely to be stillborn than girl babies. I thought it common knowledge.
For example: -
"Males had a 48 per cent increased risk for stillbirth after adjusting for birth weight, gestation and maternal diabetes," the authors wrote in the Australian and New Zealand Journal of Obstetrics and Gynaecology.
http://www.smh.com.au/lifestyle/life/boys-more-likely-to-be-stillborn-20090407-9yrm.html
If a pregnant woman has poor health, she is more likely to have a stillborn baby, and that baby is more likely to be a boy.
Again, I really don’t know how the author forgot to mention it.
Rob Brooks
Rob Brooks is a Friend of The Conversation.
Professor of Evolutionary Ecology; Director, Evolution & Ecology Research Centre at University of New South Wales
A very intriguing suggestion, Daniel. A temporary drop in the infanticide rates (which are both well-documented and implicit in the heavy male-biased ratios) might well cause an effect of this sort. I'm not sure if the survey method would allow Song to test this idea though, but other data sources might.
Infanticide and stillbirth are not necessarily alternatives to the T-W adaptive hypothesis, as we expect many mechanisms may underpin T-W effects in humans. I argue in Sex, Genes & Rock 'n' Roll that the plague of female-biased infanticide, abortion and neglect may stem in part from T-W effects.
Thanks for the suggestion.
Dale Bloom
Analyst
Rob Brooks,
Although your theory of infanticide doesn't match up with the data regarding stillborn babies, with a significantly higher number of male foetuses being stillborn.
"A national analysis of more than 16,000 births has revealed male foetuses have a significantly higher risk of being stillborn, as do Aboriginal babies and those with mothers with type one diabetes."
Perhaps your theory (which I personally wouldn’t regard as being a theory) has to be given more thought.
Bronwen Dalton
Senior Lecturer, School of Management at University of Technology, Sydney
A fascinating article. I note that the image above is of a woman in traditional Korean dress (although the small girl is in Chinese clothes). I read to see if you reported what has happened to sex ratios in North Korea since the famine in the mid 1990s when it is estimated that between 600,000 and one million people, or about 3–5 per cent of the pre-crisis population, died. In the years since one interesting development is that there may be a decline in a preference for sons. Based on calculating…
Read moreDale Bloom
Analyst
The world wide figure for baby boys to baby girls is about 105 boys born to 100 girls. It has been that way for centuries, and it doesn’t matter too much what Nth Korea is doing, because the so-called theory is about baby boys being born in every country.
The whole theory has a cental belief that boys require more resources to raise than girls, but I have never seen any data that supports that central part of the theory.
The theory has more holes than a piece of Swiss cheese after mice have been feasting on it.
Rob Brooks
Rob Brooks is a Friend of The Conversation.
Professor of Evolutionary Ecology; Director, Evolution & Ecology Research Centre at University of New South Wales
Thanks for that, Bronwen. Very helpful insight and I'll dig up that paper when I get back to work in a week or two.
South Korea has also experienced a big turn away from female infanticide and male-biased care, but that seems linked to educational initiatives from the S Korean government.