Steve Reuland posted Entry 611 on November 11, 2004 01:30 PM.
Trackback URL: http://www.pandasthumb.org/cgi-bin/mt/mt-tb.fcgi/609

There have long been attempts by evolutionary biologists and evolutionary psychologists to understand just what effect different mating systems have on evolution.  Certainly, mating systems in which a male is able to sire many offspring with many different females will cause natural selection to favor different traits than one in which a male is limited to one female.  Additionally, a mating system that causes some types of competition to become more extreme can greatly reduce other types of competition.  Gorillas, for example, are polygamous, with one male controlling a harem of females to which he has exclusive access.  In a situation such as this, there is extreme competition for gaining control of such a harem, but little competition for mating once dominance is established.  As a result, there is selective pressure for gorilla males to beat back other males and become dominant, and the results are obvious:  gorillas exhibit the greatest sexual dimorphism of any living primate, with males almost twice as large as females.  Selection favors the big and brawny types who can successfully fight off their rivals.  But there is another form of competition, one in which gorillas aren’t subjected much to — competition between sperm to be the first to reach the egg.  There are any number of ways in which sperm from different males can compete, the most obvious of which is through simple quantity.  But since male gorillas can be pretty well assured that the females in their harem will mate only with themselves, their sperm has no competition, and it shows when it comes to quantity:  male gorillas have the smallest testes to body weight ratio of the great apes.  That’s right, the burly boys of the ape world aren’t really packing that much down below, but I still wouldn’t poke fun at them. 

You can see how mating systems affect evolution throughout the primate world of which we are a part.  Chimpanzees, whose females are very promiscuous, tend to have little sexual dimorphism but a very large testes to body weight ratio.  For them, the selective pressure is more heavily focused on post-copulatory sperm competition rather than simple fighting ability.  Gibbons, on the other hand, are strictly monogamous.  They have little sexual dimorphism and a small testes to body weight ratio.  For them, there isn’t much competition to fight off rival males or to thwart rival sperm.  We humans seem to be somewhat in the middle, with a moderate amount of sexual dimorphism and a moderate testes to body weight ratio, indicating that during our evolutionary past, we weren’t nearly as monogamous as we’d like to think.  But at least we weren’t as slutty as the chimpanzees.  (Our lack of total monogamy is corroborated by additional evidence from our genome, some of which Carl Zimmer talks about here.)

Now a new piece of the puzzle has been tossed into the mix, thanks to some new research appearing in Nature Genetics by Steve Dorus and coworkers.  The new study, titled “Rate of molecular evolution of the seminal protein gene SEMG2 correlates with levels of female promiscuity” (subscription required), focuses on another facet of sperm competition other than simple quantity.  In this case, post-copulatory semen coagulation.

The gene that the authors looked at codes for the protein semenogelin, which undergoes covalent cross-linking to form a seminal coagulum after ejaculation.  A protease later breaks up the coagulum and releases the sperm.  It’s been proposed that this process of forming a coagulum and later releasing the sperm aids in getting sperm to where they need to go, and, more importantly for our discussion, preventing a rival’s sperm from getting there first.  For one thing, the semen coagulum can block other semen from getting to its destination, in some cases forming a “plug” which blocks access altogether.  A previous study showed that there is a strong correlation between the thickness of the coagulum and the promiscuity of females.  Species in which females mate with multiple males have thicker or more plug-like coagula. 

Dorus and coworkers decided to look at the evolution of semenogelin in humans and several related species, and attempted to see if there was a correlation between evolutionary rate and mating system.  What they found was that in species in which females mate with multiple males, semenogelin has been evolving faster than in those who are monogamous or polygynous.  The results are shown in graph A of the following figure, and they look quite similar to the correlation between female promiscuity and testes size shown in graph B. 

http://www.pandasthumb.org/pt-archives/ng1471-F2.gif

On the Y-axis, the lower-case Greek letter ‘omega’ represents the ratio of nonsynonymous to synonymous mutations.  A ratio above 1 indicates positive Darwinian selection, whereas a number less than 1 represents purifying selection, and a ratio of exactly 1 represents neutral evolution.  We can see that there appears to have been substantial positive selection in chimpanzees (who also have the highest slut factor), somewhat less but still decidedly positive for macaques, and around one or less for the rest of us prudes.  We humans, yet again, fall somewhere in the middle. 

These results suggest that post-copulatory sperm competition drives the evolution of semenogelin.  To complement this study, we’ll need to see what biochemical properties of the protein have presumably been selected for in chimpanzees, for example, and how they give sperm a tail up on the competition.  For now we don’t know much other than the fact that semenogelin has undergone positive selection, and that it’s highest in polyandrous species.  It’s interesting to note that the protease that liquefies the coagulum, kallikrein 3, did not show any signs of positive selection.  That would indicate that it’s some property of coagulum formation, rather than liquefaction, that allows some sperm to do better than others.  There’s bound to be lots more fascinating research in this area, thanks to those intrepid scientists who study monkey splooge. 
 

Update:  It’s been brought to my attention that people may lose their appetite from reading this topic, and that they should be forewarned.  Since I am a caring person, here goes:  Don’t read this post before mealtime.

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Comment #10181

Posted by Don Knotts on November 11, 2004 2:20 PM (e)

Next time, please don’t post something like this just before lunchtime Pacific time. Thanks a million.

DK

Comment #10183

Posted by Ruidh on November 11, 2004 3:52 PM (e)

If being forewarned is being forearmed, I think I’ve been backarmed.

Comment #10184

Posted by Reed A. Cartwright on November 11, 2004 4:08 PM (e)

How does one get ape ejaculate?

Comment #10186

Posted by Steve Reuland on November 11, 2004 4:19 PM (e)

Reed wrote:

How does one get ape ejaculate?

It probably involves a six pack of beer and the laterst Koko video.

Comment #10187

Posted by Jeff L. on November 11, 2004 4:19 PM (e)

I’m not sure, but I think in spurts. :>

Comment #10189

Posted by Great White Wonder on November 11, 2004 4:47 PM (e)

Ape ejaculate posts and it’s not even Friday!

It’s news to me that human females average about 1.5 male partners per periovulatory period (which is around 2 months? the month before and after ovulation?). I would have guessed that it’d be closer to 1 in that time frame.

Where did that data point come from? I remember some girls in high school and college got around but it seemed that even the flaky ones could stand making it with just one guy for two months. Are there that many swinging human females in the population at large?

Comment #10190

Posted by Steve Reuland on November 11, 2004 4:57 PM (e)

Here’s what the authors say:

Mating systems in humans are varied and continuously changing due to cultural influence. The mean number of partners is probably somewhat greater than one, though the precise mean across culture and time is difficult to assess.

This is science lingo for, “I keep hearing that there are these floozies out there that’ll do it with anyone, but for some reason I never seem to meet them.”

Comment #10191

Posted by Flint on November 11, 2004 5:11 PM (e)

I interpret the number to mean that at any given time, about half of sexually mature humans are involved in two sexual relationships. This isn’t exactly wildly promiscuous.

Comment #10192

Posted by Great White Wonder on November 11, 2004 5:18 PM (e)

I interpret the number to mean that at any given time, about half of sexually mature humans are involved in two sexual relationships. This isn’t exactly wildly promiscuous.

Maybe not. But I work in an office of 20 people and I think we’d all be shocked if more than one of us was involved in two sexual relationships at any given time.

Whether the number is 1.1 or 1.5 wouldn’t seem to affect their conclusion. But if I was to pull a number out of my butt like they appear to have done, I probably would have gone with the lower number.

Comment #10193

Posted by Steve Reuland on November 11, 2004 5:35 PM (e)

GWW wrote:

Maybe not. But I work in an office of 20 people and I think we’d all be shocked if more than one of us was involved in two sexual relationships at any given time.

That’s probably why they’re not telling you about it.

Studies done to quantify the level of fooling around (those based on DNA evidence, not unreliable self-reporting) tend to show a shockingly high level of cuckholdry. Up to 40% in some urban areas. I don’t think the median number of partners is 1.5, but that doesn’t sound too unreasonable as a mean when you take porn stars into consideration. We also have to keep in mind that not all cultures are like ours.

Comment #10194

Posted by Reed A. Cartwright on November 11, 2004 6:40 PM (e)

Cuckholdry studies in humans are always underestimates because once they start finding cuckholdry, women begin to drop out of the studies.

Comment #10196

Posted by Great White Wonder on November 11, 2004 7:04 PM (e)

“I don’t think the median number of partners is 1.5, but that doesn’t sound too unreasonable as a mean when you take porn stars into consideration.”

Is it reasonable to accept porn stars into the sample? I’m not sure which movies you’ve seen recently, but my impression is that very little ejaculate finds its way into the vaginas of female porn stars.

“We also have to keep in mind that not all cultures are like ours.”

But most cultures are monogomous and a substantial fraction of women in most cultures are not having sex with any partner in a two month period.

I’m still not convinced the mean number of partners per woman is much greater than 1. It may even be less than one. And that includes prostitutes (who may be subject to a disclaimer similar to that of porn stars).

Comment #10199

Posted by Flint on November 11, 2004 9:48 PM (e)

I would expect the usual 80-20 rule to apply in most cultures, where 80% are monogamous or celibate during any given 2 month period, while 20% are out playing the field. I’ve known quite a few men who hit on every women they come near, just on general principles. It works more often than I would have expected, had I not seen it happen.

But as far as I know, humans are the only species on those graphs that makes a concerted effort to conceal their sexual activity. My gut feeling is that the 1.5 figure isn’t at all unreasonable. If GWW feels deprived, I point out that prices vary widely…

Comment #10200

Posted by Steve Reuland on November 11, 2004 9:53 PM (e)

I’m joking about the porn stars, obviously, but my point is that the number of sex partners is not distributed equally. It may be that “normal” women have only one partner, but as long as there are some women who have many, then the average will be greater than one.

As for cutural differences, yes, most cultures have some form of ritualized monogamy, but adultry is common just about everywhere. And in some places, it’s common for women to have children at a young age before they’ve entered into a socially recognized life-long monogamous relationship, and it’s anyone’s guess as to who the father is. That happens quite frequently in far off exotic places like California or Mississippi.

Comment #10202

Posted by jay boilswater on November 12, 2004 1:57 AM (e)

would this be considered a seminal advance in the study of early hominid dimorphism?

Comment #10209

Posted by gaebolga on November 12, 2004 10:47 AM (e)

Jay’s post is semen more and more pun-ish with every read….

Comment #10259

Posted by Karl Lembke on November 13, 2004 7:03 PM (e)

Discussions of human polygamy should also take into account “serial monogamy”, in which people will divorce and re-marry.

(It looks like people have gone ape-s**t over this topic.)

Comment #10309

Posted by AndyS on November 15, 2004 1:47 AM (e)

Man, you biologists must really have a way with the ladies!

Anyway, the ideas here about human males and females fits with my view of the world (expect my own conclusion about biologists being hot dates - that comes as a surprise).