Category: evolution

Nothing in Biology Makes Sense: Searching for Ronald Fisher

Nothing in Biology Makes Sense: Searching for Ronald Fisher

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Geneticist Ronald A. Fisher. Photo via WikiMedia Commons.

This week at the collaborative science blog Nothing in Biology Makes Sense!, my lab-mate John Stanton-Geddes writes about the current state of evolutionary genetics, as presented at the recent Evolution meetings in Ottawa:

One theme that emerged through the meeting was “The genetic basis for [insert trait here]. While this goal of mapping phenotype to genotype has been a primary goal of many evolutionary ecologists since the first QTL mapping studies, it has recently come under strong criticism, notably in a fantastic paper by Matthew Rockman in the journal Evolution last year, but also by Pritchard and Di Rienzo 2010 and in a forthcoming article by Ruth Shaw (full disclosure: Ruth was my PhD advisor) and Mike Travisano.

Readers of Denim and Tweed will recognize that John’s complaint about our ongoing fixation (ha!) on individual genes of large effect mirrors some of my own recent thinking. So naturally, I think you should go read the whole thing.◼

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Is female orgasm adaptive? Let’s ask the clitoris.

Is female orgasm adaptive? Let’s ask the clitoris.

ResearchBlogging.orgWhether or not a trait is an adaptation, shaped by natural selection for a specific function, can be a surprisingly contentious question in evolutionary biology. When the trait in question belongs to human beings, though, “contentious” reaches a whole new level—because when evolutionary biologists consider humans, their conclusions get personal.
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Erotic sculpture on temple wall, Khajuraho, India. Photo by Abhishek Singh aka Bailoo.

Among the myriad traits and behaviors of Homo sapiens evolutionary biologists might choose to study, few can be as personal as the female orgasm. The adaptive function of male orgasm is about as clear-cut as possible—it’s a mechanistic necessity for uniting a sperm with an egg. But while female orgasm is enjoyable (or so I am told; this is as lousy a point as any to admit that my expertise in this phenomenon is purely academic), it isn’t necessary for fertilization. No man can be a father without having had at least one orgasm, but a woman could conceivably give birth to a huge family without having any.

To explain the existence of female orgasm in an evolutionary context, then, biologists have two options: (1) discover a way in which female orgasm shapes reproductive success indirectly, or (2) conclude that female orgasm isn’t an adaptation. Possibilities advanced for the first option range from the benefits of closer bonding with a mate—sex is, after all, about more than mere reproduction—to suppositions that the contractions associated with orgasm help draw semen into a woman’s reproductive tract.

The argument in support of non-adaptive female orgasm takes a developmental perspective: that female orgasm is really male orgasm, as experienced in a female developmental context. That is, women have orgasms for the same reason men have nipples—because the anatomies of both sexes are constrained by their origins in the same underlying developmental program. If this is the case, natural selection would work to optimize male orgasm, without necessarily affecting female orgasm—and that suggests a way to test whether female orgasm is an adaptation.

Natural selection removes less-fit versions of traits from a population—making that trait less variable within the population under selection. Traits that don’t affect survival or reproductive success, on the other hand, are free to accumulate variation via mutation. So non-adaptive traits can be identified by comparing their variation to traits with known adaptive functions.

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Who cares what natural selection thinks, anyway? Photo by

Psychologist Kim Wallen and philosopher of science Elisabeth Lloyd (who had advanced the hypothesis that female orgasm is non-adaptive in a 2005 book) made just such a comparison in a 2008 study. Variation in female orgasm would be challenging to measure, so they used the clitoris as an anatomic proxy. This let them use the penis—which shares a developmental origin with the clitoris and is presumably under natural selection associated with male sexual function—as an adaptive standard for comparison. In comparison to (flaccid) penis length, Wallen and Lloyd found that clitoris length was indeed more variable [$a]. As a second control, the authors also compared variation in clitoris and penis length to variation in the length of women’s vaginas, understanding that this trait, unlike the clitoris, is important for female reproductive success. Vaginal length turned out to be about as variable as penis length, and much less so than clitoris length.

There are several objections to be made to Wallen and Lloyd’s analysis, and many were made in a response [$a] by evolutionary biologist Vincent Lynch. Lynch objected to the use of length as the focal measure for the size of the clitoris, and showed that clitoral volume was about as variable as penile volume. (I would add that the study of social insects Wallen and Lloyd cite as a precedent for their analysis isn’t actually focused on variation, but on the symmetry of traits under consideration, which is not quite the same thing.) More critically, though, Lynch points out that there isn’t any known relationship between clitoral size and ability to achieve orgasm—so the data don’t have the bearing on the question that Wallen and Lloyd assigned in the first place. Lynch concluded that female orgasm is an adaptation after all—a more conservative interpretation of his result is that we can’t answer the question by measuring clitorises.

Understanding the evolution of human sexual behaviors can help us to figure out how best to navigate the tricky business of a sexual relationship with another person—an approach most recently exemplified in the book Sex at Dawn. But we also tend to view evidence that natural selection favors a particular trait or behavior as a kind of approval, or as evidence of what is “natural.” That’s silly. Whether or not they help to make more babies, orgasms are fun, and they’re a wonderful part of our most intimate expression of affection and love. In some respects, that’s all we need to know.


Crespi, B., & Vanderkist, B. (1997). Fluctuating asymmetry in vestigial and functional traits of a haplodiploid insect. Heredity, 79 (6), 624-30 DOI: 10.1038/hdy.1997.208

Lynch, V. (2008). Clitoral and penile size variability are not significantly different: lack of evidence for the byproduct theory of the female orgasm. Evolution & Development, 10 (4), 396-7 DOI: 10.1111/j.1525-142X.2008.00248.x

Wallen K, & Lloyd EA (2008). Clitoral variability compared with penile variability supports nonadaptation of female orgasm. Evolution & development, 10 (1), 1-2 DOI: 10.1111/j.1525-142X.2007.00207.x

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