Tuesday, May 17th 2011
Sons and daughters and differential parental investment
One of my favorite rhetorical tricks is asking my students a question that has an obvious answer based on cultural expectations, but is wrong. So every year, when I start to teach my students about parental investment, I ask:
Who is harder to raise, sons or daughters?
I’ve asked by a show of hands and with iClickers, over the years, and the room of 750 is almost unanimous: daughters are harder to raise. So, then I get off the stage and walk around a bit. What do you mean by that? I ask.
Girls cause more gray hairs.
Girls cause more trouble when they start to like boys.
Girls are more work, and cost more money, since they shop all the time.
Girls talk back more.
And of course, there is always the saying that girls steal some of their mother’s beauty.
So then I show them this:
|From Helle et al 2002.
Here is a graph of maternal longevity based on the number of sons or daughters they have. This data was based on a historical population from Finland from 1640-1870 using church records (Helle et al 2002). As you can see, the more sons mothers bear, the shorter their lifespans. You see the opposite for daughters. So sons have a negative impact, and daughters have a positive impact. This same trend has been found in records from a Flemish village (van de Putte et al 2003, 2004), where sons negatively impact lifespan but not daughters. Interestingly, data from church records from the field site where I work in rural Poland provides a slightly different picture: every offspring of either sex reduced lifespan by about 95 weeks (Jasienska et al 2006).
|From Jasienska et al 2006.
Once students see these graphs, they quickly realize what is going on. Generally speaking, girls help mothers more at home in terms of chores and alloparenting. And in many cultures, particularly the historical ones studied so far, sons are costly because parents invest more in them, to help launch their own families. Daughters, not so much. In the Polish population, there may be other factors where daughter investment is important, or it is just costly to have so many offspring and you have maternal depletion regardless of daughter help.
Sex bias in parental investment is an important part of understanding both the biology and culture of parenting, and the developmental trajectories of children. The Trivers-Willard hypothesis, which has been tested many times in humans and animals, suggests that parents should invest more in sons when conditions are good, and more in daughters when conditions are bad. That is, when you have lots of resource you should put it towards a son in order to increase the chances he will have high reproductive success, since his is assumed to be more variable and high effort could lead to high reward. But in periods of low resource, daughters are a good bet because they are more likely to have at least some reproductive success no matter what.
|From Hrdy 1990.
Of course, differential investment based on resources is further conflated in humans due to culture and, I would contend, our almost universal favor for patriarchy (Smuts 1995). Here is an image of an Indian family waiting at a clinic. There is a mother, an older son, and a twin boy and girl. Notice the extreme difference in health between the infant boy and girl – they are twins, yet the infant girl is emaciated. This is because in this population sons are always fed and cared for first, and whatever is left over, if there’s anything, is given to daughters.
So, parental investment can have real effects on the parent in terms of lifespan, and perhaps also their own future reproductive capabilities. Further, the conditions under which you may have children can vary, but how much a parent chooses to invest in their children varies too.
The piece of this that may be toughest to parse out, particularly in humans, is how the condition of the mother (or parents) can vary, and how that variation impacts the sex ratio of their children. In some species, like red deer, it is easier to imagine a mechanism: these animals have diapause, a period where their embryos are dormant until it is a good time to gestate and bear them. It is easier to insert some kind of selection process into a period where several embryos are all “frozen” and sex has been determined. But what about humans that produce singletons and invest huge, overlapping amounts of support to their children over decades? How would a sex bias based on maternal condition operate? And is there anything the offspring can do about it?
Changes in maternal breast size during pregnancy
It turns out that measurements as easy as stepping on a scale, and knowing your bra size, can begin to unpack the answer. First, a confession: I consider the author of this paper Andrzej Galbarczyk more than a colleague, but a friend. Andrzej is the graduate student who oversaw my Polish field site last season (Mogielica Human Ecology Study Site, director Dr. Grazyna Jasienska). He has translated consent forms and surveys for me and we’ve had many valuable and important conversations about my fieldwork. He is a smart, kind and thoughtful person and scholar. So, I let him see an early draft of this post to make sure I understood his point of view.
Galbarczyk performed an internet survey in Poland with 120 women, where he asked them to report their pre-pregnancy weight and bra size, their bra size directly after giving birth, and the sex of their offspring. He found two notable differences in these women: mothers of daughters weighed less before pregnancy, but had a greater changes in breast size during pregnancy.
The evidence about maternal pre-pregnancy weight is consistent with the Trivers-Willard hypothesis, as mothers who had sons were more likely to be heavier, and thus have more resource to invest. The second significant difference, that mothers with daughters had larger breasts after pregnancy, seems could be argued either way: Galbarczyk argues that it supports Trivers-Willard because mothers of sons could have been devoting more resource to growing their offspring rather than their breasts.
In other animals and primates particularly, mothers of male infants produce more energy-dense milk, yet mothers of female infants may produce a greater quantity of milk (Hinde 2009). And breast size is a pretty noisy signal of milk quality or quantity. So, what is the meaning of this difference in breast size?
Adaptation or physiological inevitability?
Galbarczyk suggests the difference is related to the evolutionary underpinnings of human female breasts. Women develop breasts around puberty, and though they certainly change in size and shape over time, keep them their whole lives. Other animals develop their mammary glands only shortly before lactating and then they regress again. Many contend that human breasts are an honest signal of fertility. This is at least partially confirmed by the correlation between breast size and estradiol concentrations (Jasienska et al 2006).
Galbarczyk thinks that the larger breasts seen in postpartum mothers to daughters may be a way to attract a mate for parental care. Perhaps this would help where she has given birth to the less-favored sex and needs to really convince him to participate; this could be a signal from the mother or the female fetus. Or maybe by appearing more attractive, she can have another reproductive opportunity, which would give her a chance to have a son.
You all know how I feel about evolutionary storytelling. In certain ways I do find this particular argument compelling, from the perspective of the Trivers-Willard hypothesis. But the evidence for the adaptive scenario around breast size is circumstantial.
Also, I don’t want this story to detract from some very interesting data: remember that Galbarczyk found that in this population, mothers of daughters weigh less before pregnancy, and develop larger breasts afterwards. Very cool. So perhaps we should consider a mechanistic, rather than adaptive explanation?
I have two thoughts about this, both related to androgens (androgens are the class of hormone that testosterone falls under). First, I wonder if there is an effect of fetal androgens from a male fetus on breast size. If so, mothers of daughters would have larger breasts simply because they aren’t having their breast tissue growth or density suppressed by androgens. It could simply be physiology that doesn’t have adaptive meaning.
Second, the mothers of sons were heavier before pregnancy. Heavier individuals tend to have higher circulating insulin levels, and the ovary can respond to higher insulin by producing more androgens (Poretsky 1991, Dimitrakakis et al 2004). So you could have a suppressive effect on breast size from that avenue as well. You don’t need an adaptive scenario for either of these mechanisms, just a consequence of how hormones work.
I would love to see Galbarczyk or someone else follow up on these thought-provoking results by measuring women, rather than relying on self-report, and by measuring their estradiol, progesterone and androgens. Understanding the different factors and motivations that lead to sex differential investment and outcome is a great field of study, and this work gets us thinking in a new direction.
Dimitrakakis C, Jones RA, Liu A, & Bondy CA (2004). Breast cancer incidence in postmenopausal women using testosterone in addition to usual hormone therapy. Menopause (New York, N.Y.), 11 (5), 531-5 PMID: 15356405
Galbarczyk A (2011). Unexpected changes in maternal breast size during pregnancy in relation to infant sex: An evolutionary interpretation. American journal of human biology : the official journal of the Human Biology Council PMID: 21544894
Helle, S. (2002). Sons Reduced Maternal Longevity in Preindustrial Humans Science, 296 (5570), 1085-1085 DOI: 10.1126/science.1070106
Hinde K (2009). Richer milk for sons but more milk for daughters: Sex-biased investment during lactation varies with maternal life history in rhesus macaques. American journal of human biology : the official journal of the Human Biology Council, 21 (4), 512-9 PMID: 19384860
Hrdy, S. (1990). Sex bias in nature and in history: A late 1980s reexamination of the “biological origins” argument American Journal of Physical Anthropology, 33 (S11), 25-37 DOI: 10.1002/ajpa.1330330504
Jasienska G, Nenko I, & Jasienski M (2006). Daughters increase longevity of fathers, but daughters and sons equally reduce longevity of mothers. American journal of human biology : the official journal of the Human Biology Council, 18 (3), 422-5 PMID: 16634019
Poretsky L, Seto-Young D, Shrestha A, Dhillon S, Mirjany M, Liu HC, Yih MC, & Rosenwaks Z (2001). Phosphatidyl-inositol-3 kinase-independent insulin action pathway(s) in the human ovary. The Journal of clinical endocrinology and metabolism, 86 (7), 3115-9 PMID: 11443175
Friday, March 4th 2011
|Figure 1. My apologies to Baby Jaguar
for not finding a picture that included
My daughter will be three in just a few weeks. She loves telling stories. These stories have the same, uncomplicated arc every time: she and her friends Dora, Diego, Boots and Baby Jaguar go on an adventure to rescue Mommy from the giant condor. Or sometimes Mommy and Dora and Diego and Boots and Baby Jaguar are rescuing her. Or sometimes Daddy does the rescuing.
There is almost always a net, then a pair of Rescue Scissors needed to cut the captive free. But the variation in these stories is very small, the framework borrowed heavily from one of the few mythologies known to my little girl: Dora the Explorer.
Evolutionary psychology is often a kind of story-telling, and instead of borrowing from a preschool cartoon they borrow from the concept of anisogamy. Anisogamy is sexual reproduction formed by unequal gametes, in our lineage a big egg made by females and little sperm made by males. This provides the foundation for differential reproductive investment, where females often put in the time and effort of gestation, lactation and care. From here, proponents of EP see essential differences between what men and women want in relationships, and the kinds of relationships that are optimal, and a model this broad makes it possible to shoehorn any behavior into its adaptive framework.
|Figure 2. The actual image that
accompanied Tierney’s column.
Enter John Tierney, my (not) favorite journalist for the New York Times. This is the man who thinks that sexism is a radical act (I am referring to his charming articles on gender disparities in science). So I suppose I shouldn’t have been surprised when he outed himself as an EP fanboi in his most recent piece, “The Threatening Scent of Fertile Women.”
Tierney covered the work of Jon Maner and others who have studied relationship maintenance – the suite of behaviors that keeps a couple together. In particular, Tierney focuses on the problem of the wandering eye, or rather, the possible mechanisms that prevent it in a monogamous couple. The idea here is that relationship maintenance is evolutionarily adaptive, because when a couple stays together it is easier to raise offspring and increase reproductive success.
The range of explanations
The study that frames Tierney’s column is Miller and Maner (2010). Thirty eight undergraduate men rated the attractiveness of a woman with whom they interact, at several points over her menstrual cycle. The authors found NO relationship between where a woman is in her cycle and how attractive a single man finds her, but a negative relationship between the chance a woman is fertile and how attractive a partnered man finds her.
What do Miller and Maner (2010) discuss, and what is the idea Tierney is so enamored with?
“It’s possible that some of the men in Florida were just trying to look virtuous by downgrading the woman’s attractiveness, the way a husband will instantly dismiss any woman pointed out by his wife. (That Victoria’s Secret model? Ugh! A skeleton with silicone.) But Jon Maner, a co-author of the study, says that’s unlikely because the men filled out their answers in private and didn’t expect the ratings to be seen by anyone except the researchers.
“It seems the men were truly trying to ward off any temptation they felt toward the ovulating woman,” said Dr. Maner, who did the work with Saul Miller, a fellow psychologist at Florida State. “They were trying to convince themselves that she was undesirable. I suspect some men really came to believe what they said. Others might still have felt the undercurrent of their forbidden desire, but I bet just voicing their lack of attraction helped them suppress it.””
This conjecture is unconnected to the study’s methodology and results. Nowhere in that study did they assess the participants’ state of mind or ask them how they felt about this. How do we know they were trying to convince themselves of anything? This finding, while interesting, does not test their hypothesis for an evolutionary framework for relationship maintenance that includes adaptively suppressing attraction to others.
Maner et al (2009) studied the attention people pay to images of attractive people of the opposite sex when first exposed to sexual words like “lust” and “kiss.” They recruited 120 straight undergraduates, thirty six of whom were in committed relationships. Individuals in committed relationships paid far less attention to the attractive images than those not in relationships. Tierney titters,
“The subliminal priming with words related to sex apparently activated some unconscious protective mechanism: Tempt me not! I see nothing! I see nothing!”
I’ve done my own share of human subjects research, and subjects will often tell you or do what they think you want, or they will just not be honest if they don’t want you to know the truth. What if, as originally posed by Tierney himself, the respondents weren’t warding off temptation but wanted to look virtuous? What if, now bear with me because this might seem crazy, the people in these studies were in love with their partners and genuinely uninterested in anyone else? Too often EP wants to provide a single explanation for a behavior, when the range of possible explanations far exceeds their hypothesis.
An anthropological perspective
Jamie Jones, Associate Professor at Stanford and blogger at Monkey’s Uncle describes anthropology like this,
“…[A]nthropology is the science charged with explaining the origin and maintenance of human diversity in all its forms. To achieve this end, anthropology must be unapologetically grand in its scope. How can we explain human diversity without documenting its full extent, through both time and space, and across cultures? … Where does the tapestry of human diversity come from and how is it that we continually manage to resist powerful homogenizing forces and hang on to our diversity? What commonalities transcend local difference to unite all humanity? How is it that civilizations rise and fall? And what is the fate of humanity?”
Jamie beautifully depicts the importance of documenting and understanding diversity even in the face of efforts to simplify human nature. Thus, to me, an anthropological perspective is often at odds with EP explanations for behavior.
An anthropological perspective asks, what happens if you take these basic observations and, instead of deciding on a favorite explanation and applying it to everyone, put them into a model in which you can vary context (age/sex specific mortality rates, distribution of resources, what have you) and see what range of strategies actually give fitness benefits? That is, when you actually throw some variation into the equation, is this still the best strategy for the partnered men with whom Tierney feels simpatico?
Right now we don’t know. Much psychological empiricism rests on undergraduates who participate in studies for course credit. When one wants to make connections to evolutionary adaptedness, they may be a place to start, but not end.
I have a real problem with continuing to use this population to make statements of universality for all humans. Undergraduates usually are trying to avoid pregnancy and build their financial and social capital, so relationship maintenance for the sake of reproductive success rarely exists. Until we can show that relationship maintenance, and the particular behaviors Miller, Maner and others study within that are shown across many populations, and particularly across reproductively-aged folks, their argument for adaptation fails.
|Figure 3. Celebrations of marriage.
Another problem is that most work on relationships in EP tends to be heteronormative, meaning that they think nothing of assuming that either everyone is straight, or the universally best behavioral strategy is to be straight. They also tend to assume that the best strategy is to be monogamous, with occasional sneaky infidelity permitted if one can get better genes or more offspring that way (keep in mind that there is a difference between what might be biologically advantageous in a certain context, and what is culturally appropriate – the argument here is not against the culture of monogamy).
But heterosexual monogamy is only one reproductive strategy of many that humans employ. Depending on how you measure it, monogamy and polygyny (single male, multi female marriage) vie for the most frequent strategy – in fact, polygyny occurs in about 80% of modern human societies (Murdock and White 1969). There are even a few rare populations that practice polyandry, which is the marriage of a single female and multiple males. And, even in those populations where monogamy is practiced, serial monogamy is far more frequent than lifetime monogamy: this means that individuals have a series of monogamous relationships rather than find one mate for life (so no, divorce is not a modern human invention).
When taking an even broader, comparative perspective, monogamy isn’t practiced by our closest relatives at all. Chimpanzees and bonobos, both equally related to us, are promiscuous. This is a scientific term for a reproductive strategy that involves females and males making reproductive decisions to mate with many individuals at each fertile period. Bonobos are also promiscuous, but they also use heterosexual and homosexual sex to reduce stress and aggression, and form bonds among one another. Gorillas, our next closest relative, are polygynous. Orangutans are very solitary, but essentially promiscuous. It’s only once you delve into the lesser apes, the gibbons, that you see any monogamy, and they are far less monogamous than we first thought (Brockelman et al 1998).
Maintaining a heterosexual, monogamous relationship is certainly advantageous at certain times, in certain contexts. But it is not universally adaptive, even within humans. Without anyone studying these behaviors in populations that use different reproductive strategies, and in the absence of comparative data to support these assertions, we are at an impasse.
In the words of a friend, EP is plugged into evolutionary theory with little more than a ratty old extension cord. EP takes some very basic, ancestral conditions, like differential costs of reproduction, and uses it in a sufficiently vague way that any behavior can relate to females generally being the ones to put in all the time and effort into making babies. Yet EP often ignores the three conditions necessary for natural selection, the mechanism for evolution. For natural selection to act on a trait, the trait must be variable, heritable, and produce differential reproductive success. Rarely does EP understand variation in a trait, rarely does it examine whether said trait has a genetic component, and rarely does it test whether their trait confers a reproductive advantage.
Are fertile women a threat to partnered harmony, their scents providing a temptation that noble men must suppress? I can’t rule it out, but I also think it is one of the least likely of many possible explanations.
Unfortunately for readers of the New York Times, Tierney loved this idea more than he loved interrogating it.
I’d like to thank Charles Roseman, friend, faculty curmudgeon and Bastard Colleague from Hell, for taking a look at an early draft of this post and providing commentary crucial to its improvement. Any rhetorical or scientific errors are my own.
Brockelman, W., Reichard, U., Treesucon, U., & Raemaekers, J. (1998). Dispersal, pair formation and social structure in gibbons ( Hylobates lar ) Behavioral Ecology and Sociobiology, 42 (5), 329-339 DOI: 10.1007/s002650050445
MANER, J., GAILLIOT, M., & MILLER, S. (2009). The implicit cognition of relationship maintenance: Inattention to attractive alternatives Journal of Experimental Social Psychology, 45 (1), 174-179 DOI: 10.1016/j.jesp.2008.08.002
Miller, S, & Maner, J (2010). Evolution and relationship maintenance: Fertility cues lead committed men to devalue relationship alternatives Journal of Experimental Social Psychology, 46, 1081-1084
Murdock, G., & White, D. (1969). Standard Cross-Cultural Sample Ethnology, 8 (4) DOI: 10.2307/3772907
Dora picture: http://www.doratheexplorertvshow.com/dora/dora-explora-pics.htm
Lady magnet: http://www.nytimes.com/2011/02/22/science/22tier.html?_r=2&ref=johntierney
Same-sex marriage: http://markusisthedrug.onsugar.com/date/2009/05/07