Sexual selection (Robert Quinlan, ANTH 468 Washington State U.)

 

If you look at this skull in figure 1, what would you conclude about this animal’s diet? 

 

Figure 1. http://www.boneclones.com/BC-036.htm

 

 

You might guess that it was a meat eater, right?  It does have huge canine teeth that would appear to be well adapted to catching and eating game.  Well, if you guessed it’s a meat eater you would be wrong. This animal subsists almost entirely on shoots and leaves.

 

Now look at the skull in figure 2 below. If I told you that it was about in proportion to the skull in figure 1 (that is it’s about 80% smaller) and that it’s a full grown animal, would you think that it’s the same species as figure 1?  I bet many of you would not, but you’d be wrong. It is the same species: They’re both gorillas. The big one in figure 1 is an adult male, and figure 2 is an adult female. Such sex differences are called sexual dimorphism.

 

Figure 2. http://www.boneclones.com/BC-035.htm

 

 

Our question is then, how is that male and female gorillas are so different? They’re so different that you might not even guess, just by looking at them, that they’re the same species.  How could evolution do that to a species?  Well, we have to back to the basics of sexual reproduction to understand the selective pressure that result in often big differences between males and females.

 

Sex differences in cell division: Anisogamy

 

If you remember back to high school biology, you may recall that there are two types of cell division. One type, called mitosis, is involved in the growth and maintenance of tissues. In mitosis the cells in your body replicate their DNA and organelles, and from one cell you end up with two identical daughter cells.  The daughter cells have the full compliment of your DNA in 23 pairs of chromosomes. Your body is undergoing mitosis right now as your skin cells replicate themselves, die and slough off. For our purpose it’s no bid deal and there aren’t sex differences in mitosis that we need to be concerned about.

 

There is another form of cell division called meiosis involved in the production of sex cells.  Meiosis is considerably different that mitosis because you end up with daughter cells that only contain half the DNA of the parent cell. It works like this: First, you start with a diploid cell (with chromosome pairs) either in the testis if you’re male or ovaries if you’re female (see figure 3, which only has one chromosome pair for ease of presentation). 

 

Figure 3. http://genetics.gsk.com/graphics/meiosis-big.gif

 

That cell duplicates its DNA; then the chromosome pairs get pulled apart in the first division. In the second division the duplicated half of the chromosome pairs are pulled apart and you end up with four haploid daughter cells (they do not have chromosome pairs).  Basically you could draw a tail on the four daughter cells in figure 3 and call them sperm.  But the process is a bit different females (figure 4).

 

 

 

Figure 4. http://faculty.southwest.tn.edu/rburkett/A&P2_reproductive_system_lab.htm

 

In females there is a big difference:  At each cell division in meiosis all of the cytoplasm and organelles (junk and functional stuff) in the gonad cell go into one relatively large daughter cell.  The leftover DNA in the other daughter cell becomes a thing called a polar body, which is ultimately absorbed.  The result is that females tend to produce relatively few, huge and expensive gametes called eggs, while males produce many millions of tiny, cheap gametes called sperm.  Figure 5 gives you an idea of the relative size of egg and sperm, where the giant egg is surrounded by many tiny sperm.

 

Furthermore, once a male hits spermarche (sexual maturity coinciding with the first nocturnal emission or “wet dream” in humans) he produces millions and millions of sperm throughout his life.  Sperm are so plentiful and cheap to produce that many males cast them away with glee and reckless abandon!  In contrast eggs are relatively costly: A human female is born with all the eggs she will ever produce (about 30,000 of them) which are suspended half way through the meiotic process, waiting to finish their division into gametes, and make their way down the fallopian tube once per month after menarche.  In mammals especially, this basic difference in the “cost” of producing gametes echoes throughout much of the reproductive process.

 

Figure 5. http://www.geocities.com/CapeCanaveral/Hall/2955/imagens/fertilization.jpg

 

Think about it for a moment: From the point of conception a woman is tied to her fetus for about 10 months.  After birth women are tied to infants by lactation (breastfeeding).  During exclusive breastfeeding, when a child does not eat much supplemental food, a woman puts about 670 calories per day into milk production. That’s more than a Big Mac with some fries.  In the U.S. today women only breastfeed on average for about 3 months, but the average cross-culturally is much longer. In half the worlds cultures women breastfeed for 30 months (2 ˝ years) or longer!  So, a woman’s average minimum parental investment entails about 40 months or just shy of 3 ˝ years just to get a child to the point where someone else could take care of it. Of course 2 ˝ year old children still want and need their mother or some other highly competent caregiver.  Now consider that on average, women around the world go through menarche at about 15 (the average is about 13 in the U.S. and other developed countries).  Keep in mind that many women experience “adolescent sub-fecundity,” a period after menarche when pregnancy is unlikely.  The average human female could expect to be able to get pregnant at about 16 to 18 years of age.  After about age 30 or a bit later fecundity begins to decline and it becomes more difficult to get pregnant until about age 50 when menopause occurs and a woman’s reproductive life ends.  So then, women have about 30 to 35 years in which to reproduce.  In other words a woman’s reproductive career is about 390 months; hence, a woman attempting to maximize her fertility could expect to have about 10 offspring.  In other words, on average her maximum reproductive potential is 10.

 

Now consider a man’s minimum parental investment:  If you include dinner and a movie, then it’s about one day.  Most males reach spermarche at about 15 (13 in the U.S. and other developed countries) – though they are unlikely to get lucky for several years after.  Men don’t have menopause, and they can make sperm until death at about 70+ years (in developed countries). Hence a man’s reproductive career is about 55+ years.  There are more than 20,000 days in 55 years.  Clearly a man’s maximum reproductive potential is not 20,000.  Mulai Ismail, (Sultan of Morocco 1646-1727) is reported to have had 4,000 wives and more than 650 sons (apparently he didn’t bother counting daughters), which gives him the highest reproductive success of any human.  Of course most men can’t hope to have 1000+ children, but a man in a polygynous union with three wives might well have 20 or 30 offspring. So, let’s just compare the reproductive prospects for men and women shall we:

 

Table 1.  Sex differences in reproductive potential for men and women

 

Women

Men

minimum parental investment

1200 days (40 months)

1 day

reproductive career

32.5 years

53 years

maximum reproductive potential

10

30-1000

 

 

 

 

Female choice and male-male competition

 

This big difference has several implications:  Because reproduction is so expensive for females, they should tend to be very choosy. Remember that we define fitness as “one’s genetic representation in future generations.” During each pregnancy a female human (or other great ape, which tend to have one offspring at time) is only nurturing half of her genes in her fetus. The other half come from some male.  Hence, she should be very careful about the genes she takes “on board” to mix with her own. After all, who wants to take care of some loser’s genes that might even reduce the chance that one’s own genes make it into future generations?  Females then should look for males with good genes.  But how do you tell which males have good genes?  

 

Females should prefer signs of good genes that are difficult to fake.  One idea is that males with good genes are better able to resist pathogens (like skin infections etc.).  Another idea is that honest signals of genetic quality should be energetically expensive to make. So for example, peacocks grow huge, flashy tails that can make life hard for them. This is known has handicap theory. The healthiest peacocks can make the best tails, which are hard to make. Peahens, then, choose their mates according to the quality of their tail.  Some sexually dimorphic traits like the peacock’s tail evolved through one mechanism of sexual selection known as female choice. [needs a bit more here: Zahavi & Hamilton-Zuk links].

 

Can you imagine something analogous to the peacock’s tail in humans?  Some evolutionists suggest that artistic and musical ability might be like the peacock’s tail. Others think that beards are a possible honest signal of male genetic quality.  Maybe it’s true. How might you test those hypotheses?

 

There is another dimension of female choice. Because pregnancy and lactation are energetically expensive females should want help: It takes lots of calories to make and nurse a baby, and it’s difficult to do other things, like look for food, when you’re breastfeeding.  In species (or environments) where males can help, then females should prefer males that show signs of willingness and ability to provision her and her offspring.  One study by psychologist David Buss showed that women in 37 different cultures were, in fact, attracted to men with resources that they could invest in offspring. [needs more: links?]

 

Female choice is only half the story.  If you imagine that female reproduction is very expensive and male reproduction is very cheap, then that creates a problem for males too.  That is, access to willing females is limited and males must compete for the attention and sexual access to females.  Males with genetically heritable traits that make them better able to compete for mates are more likely to pass on their genes and those traits that make them better competitors evolve through the second mechanism of sexual selection known as male-male competition.  In many species male-male competition gets played out in combat among males.  In that case larger males tend to gain access to females, and male-male competition leads to the evolution of sex differences in body size. For example, gorilla males are 80% larger than gorilla females, which is most likely due to competition for access to mates.  Male gorillas also have large canine teeth for fighting. Both traits – large body size and canine teeth – are said to be sexually selected (fig. 1).

 

In theory there can be male choice and female-female competition. This reversal is particularly the case when males provide extensive parental care.  More accurately we should refer to inter-sexual choice and intra-sexual competition as the two mechanisms of sexual selection or mechanisms by which sex differences evolve.


What about humans?

 

Among humans we might expect some sexually selected differences.  Human females face many of the same problems that other animals face in terms of mate choice.  Women should be choosy about the genes that they bring “onboard” to mix with their own because pregnancy, lactation and later childcare are especially expensive for humans.  Mistakes in mate choice can be very costly for women.

 

Human males are able to contribute substantial parental care, either directly or by provisioning the mother and her offspring, which can free women from work that conflicts with childcare.  This fact sets up a kind of two-way street for human sexual selection: (1) Women should be choosy not only about the quality of her mate’s genes, but also about his willingness and ability to provide resources for childrearing.  (2) Men should also be choosy about their mates’ genetic quality because men may invest a lot of time and resources into a mating relationship and offspring, and men often forgo other mating opportunities to invest in one family.  Hence, men should be choosy about another quality of their mates: Their potential fidelity. After all, if man invests substantial resources in a relationship with a woman and her offspring, then he should want to be as sure as he can be that those offspring share his genes.  That means men should look for signals of fidelity along with signals of good genes.

 

Human males are about 15 to 20% larger than are human females depending on the population.  This sexual dimorphism reflects at least moderate male-male physical competition for access to mates or access to resources for mating.  There may be even greater sexually selected psychological differences between men and women.

 

In western industrial populations, like the U.S. and Western Europe we find very few psychological differences between men and women.  The differences we do see, however, are consistent with a natural history of sexual selection:  Men tend to be more aggressive than are women and this appears to be true across all cultures.  Greater male aggression is exactly what we would expect to see in a species with male-male competition for access to mates and resources for mating effort.  Sex differences in attitudes toward promiscuity might also reflect sexual selection. Women tend to have more conservative or discerning attitudes about sexual relations, which reflect greater choosiness in mate choice.  Even in today’s industrial nations women are much more likely to be left caring for children after her mate abandons the relationship than are men – a woman can’t leave her pregnant boyfriend or husband, but men can leave a pregnant mate, and many do.

 

Even with aggression and sexual “permissiveness” there is substantial overlap between men and women.  Janet Shibley Hyde conducted a “meta-analysis” of many studies of psychological sex differences, which shows that the average difference between men and woman was only about .8 to 1 standard deviation for traits that are likely sexually selected.  Figure 6 shows that there is substantial overlap between men and women for attitudes toward casual sex, for example.   In the figure the vertical lines show the average for men and women, and the bell-shaped curves show the variation around the average.  What the figure shows is that there are many women who are more accepting of casual sex than are many men, but the general tendency is for women to be less accepting of casual sexual relationships.

 

David Schmidt conducted a study of college students around the world, which shows that the variation in gender differences in attitudes toward casual sex is influenced by culture.  In societies where men and women are more equal in economic power and opportunities, there is less sex difference in attitudes toward casual sex.  But there is still some sex difference: Women still tend to be less accepting of casual affairs than are men.

 

Culture also influences what we believe to be appropriate sexual behavior for men and women.  A series of fascinating psychological studies uses as technique called the “bogus pipeline” to examine our culturally shaped notions of sexual propriety.  In the bogus pipeline method one group of subjects is given a questionnaire and is hooked up to a fake lie detector (the bogus pipeline). The other group of subjects simply fills out the questionnaire anonymously.  Terri Fisher did a bogus pipeline study of sexual experience (number of sex partners and number of one-night-stands), and the results are shown in figure 7:  Women college students tend to under-report their sexual experience, while men exaggerate their experience. 

 

Figure 6. Difference between men and women in attitudes toward casual sex. Dotted lines indicate men, solid lines indicate women

 

The bogus pipeline shows something else very important.  Notice that in the pipeline condition there is no difference in the average level of sexual experience of men and women (fig. 7 “Pipeline”).  However, there is a big difference in the standard deviation:  Men have much more variation in the level of sexual experience than do women, meaning that some men have much more sexual experience than do others.  Such variation among males is precisely what we expect to see under male-male competition and female choice – some males are being relatively excluded from the mate pool, while others are highly successful. In contrast, most women have about the same level of sexual experience.

 

Figure 7.  Expectations for appropriate gender behavior influences our responses to questionnaires.  Bars represent the mean and “whiskers” represent the standard deviation.  “Sexual Experience” includes number of partners and number of one-night-stands.

 

Summary & Conclusion

 

Basic sex differences in the costs of reproduction create different pressures on males and females.  Females should be choosy about the quality of genes they bring on board and the willingness and ability of their mate to help with childrearing.  Males should be competitive for access to choosy mates, and when men provide parental care they should be concerned about their mate’s fidelity.  Among humans these differences in costs of reproduction may account for moderate sexual dimorphism in body size, and dimorphism in aggression and acceptance of casual sex.  Cultural differences do appear to play a role in either narrowing or widening the gap between men and women.  In no society, however, have we ever observed a full reversal in sex differences:  No where are women on average larger and more aggressive than men, and no where do women appear to be more motivated on average to have casual sex than are men.